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FIELDIANA 



Zoology 

NEW SERIES, NO. 113 



Systematic Review of the Barbary Macaque, 

Macaca sylvanus (Linnaeus, 1758) 



Jack Fooden 



October 26, 2007 
Publication 1547 



PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY 



FIELDIANA 



Zoology 

NEW SERIES, NO. 113 



Systematic Review of the Barbary Macaque, 
Macaca sylvanus (Linnaeus, 1758) 



Jack Fooden 

Division of Maiiiinals 

Department of Zoology 

Field Museum of Natural History 

1400 South Lake Shore Drive 

Chicago, Illinois 60605-2496 

U.S.A. 



Accepted July 20, 2007 
Published October 26, 2007 
Publication 1547 



PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY 



© 2007 Field Museum of Natural History 

ISSN 0015-0754 

PRINTED IN THE UNITED STATES OF AMERICA 



Table of Contents 

Abstract 1 

Introduction 1 

Materials and Methods 1 

Geographic Distribution 2 

Total Population Estimates 2 

Pelage 3 

External Measurements and Proportions 4 

Caudal Vertebrae 6 

Cranial Characters 6 

Molecular Biology and Genetics 8 

DNA variation 8 

Karyology 12 

Natural History 13 

Habitats 13 

Terrestriality/Arboreality 14 

Group Size and Composition 14 

Group Fission 15 

Home Range Area and Population Density 15 

Diet 15 

Predators 17 

Intergroup Migration 18 

Reproduction 18 

Seasonality 18 

Sexual Maturation 19 

Reproductive Anatomy Notes 19 

Female Sexual Cycles 20 

Consortships 20 

Copulation 21 

Male Dominance Rank and Reproductive Success 21 

Inbreeding/Outbreeding 21 

Gestation 22 

Parturition 22 

Birth Rate 23 

Neonatal Sex Ratio 23 

Twinning 23 

Stillbirths; Infant Mortality 24 

Weaning 25 

Male-Infant Interactions 25 

Longevity; Reproductive Senescence; Death Rate 26 

Population Growth Rate 26 

Fossils 27 

Systematics 28 

Synonymy 28 

Type Series 30 

Type Locality 30 

Distribution 30 

Diagnosis 30 

Remarks 30 

Specimens Examined 31 

Evolution and Dispersal 31 

Acknowledgments 32 

Literature Cited 32 



Appendix I: Specimens Examined (Details) 42 

Appendix II: Gazetteer of XfACUA syi.iams Localities 42 

Appendix III: Centrum Length and Vertebral Count of Caudal Vertebrae in Macaca syh ams . . .52 

Appendix IV: Groip Size and Adult Sex Composition in Macaca sylvasvs Populations 53 

Appendix V: Coordinates, Geological Age, and Reported Identification of Circum-Mediterranean 
Macaque Fossils 54 

List of Illustrations 



\. Map of northwestern Africa showing extant and Holocene localities of Macaca syhanus 3 

2. Adult pelage in Macaca syhanus 

A. Male, pre-molt 5 

B. Male, molting 5 

C. Estrous female, post-molt 5 

3. Tail \estige in Macaca syhanus 7 

4. Ventral view of sacrum and caudal vertebrae in Macaca sylvanus 9 

5. Skull of adult female Macaca sylvanus 10 

6. Skull of adult male Macaca sylvanus 11 

7. Phylogenetic relationships among Moroccan and Algerian populations of Macaca sylvanus as indica- 

ted by mtDNA haplotypes 13 

8. Copulation posture in Macaca sylvanus 22 

9. Geographic distribution of circum-Mediterranean macaque fossils compared with geographic distribu- 

tion of extant populations and Holocene extinct populations of Macaca sylvanus 28 

List of Tables 



1 . Frequency distribution of elevation records of Macaca sylvanus 4 

2. External measurements and proportions in wild-collected Macaca sylvanus 6 

3. Tail length and relative tail length in Macaca sylvanus specimens collected near Azrou. Moyen 

Atlas. Morocco 8 

4. Cranial measurements and proportions in wild-collected Macaca sylvanus 12 

5. Mitochondrial DNA haplotype frequencies in Gibraltar and North African samples of Macaca sylvanus. . 12 

6. Estimated dates of divergence of Macaca sylvanus sister groups 13 

7. Habitats of Macaca sylvanus 14 

8. Frequency of terrestriality in Macaca sylvanus populations 15 

9. Home range area in natural populations of Macaca sylvanus 16 

10. Population density in Macaca sylvanus 16 

11. Number of gymnosperm and angiosperm species consumed by Macaca sylvanus populations 17 

12. Allocation of feeding time to various dietary components by Macaca sylvanus groups in Grande 

Kabylie, Algeria 18 

13. Seasonal mating periods and birth periods reported in Macaca sylvanus populations 19 

14. Mean annual birth rate in Macaca sylvanus groups 23 

15. Neonatal sex ratio in Macaca sylvanus 24 

16. Twinning frequency reported in semi-natural populations of Macaca sylvanus 24 

17. Stillbirth frequency in semi-natural populations of Macaca sylvanus 25 

18. Infant mortality rate in natural and semi-natural populations of Macaca sylvanus 25 

19. Mean annual death rate in semi-natural populations of Macaca sylvanus 27 

20. Demographic growth rate in natural and semi-natural populations of Macaca sylvanus 27 

21. Evolution and dispersal of circum-Mediterranean macaques: hypothetical chronology 31 



Systematic Review of the Barbary Macaque, Macaca 
sylvanus (Linnaeus, 1758) 



Jack Fooden 



Abstract 

The Barbary macaque, Macaca sylvanus (Linnaeus, 1758), is systematically reviewed, based 
on a study of 103 specimens and survey of relevant literature. This review includes analyses of 
external characters, cranial characters, DNA variation, and karyology. Information also is 
presented concerning natural history, reproduction, fossils, and taxonomic history. Current 
knowledge of geographic variation in M. sylvanus is insufficient to warrant recognition of 
subspecies. Although M. sylvanus is now restricted to northwestern Africa, fossil evidence 
suggests that this species is a relict of the ancestral macaque stock that originated in Africa ca. 
7-6 million years ago (Ma) and dispersed to Eurasia ca. 6-5 Ma. Macaques formerly were 
widely distributed in Europe, but became extinct there ca. 0.100-0.020 Ma; macaques have 
survived in Asia and are now represented there by ca. 20 species. An annotated gazetteer lists 
166 localities where M. sylvanus has been collected, observed, or reported. 



Introduction 



Materials and Methods 



The Barbary macaque, Macaca sylvanus, is 
unique within its genus in two obvious respects: 
(1) it is the only macaque species that is 
distributed outside of Asia, and (2) it is the only 
macaque species in which the tail is reduced to an 
inconspicuous boneless vestige or is completely 
absent. Probably as a result of its juxta- 
Mediterranean distribution and its distinctive 
tail reduction, the Barbary macaque was well 
known to ancient Phoenicians, Etruscans, Egyp- 
tians, and Greeks — including Aristotle (Mc- 
Dermott, 1938, pp. 28, 29, 38^ 88; Goudsmit & 
Brandon-Jones, 1999, p. 49). This species is the 
first primate listed in Gesner's (1551, p. 957) 
classic Historiae Aninialium, from which it was 
cited by Linnaeus (1758, p. 25). 



Division of Mammals, Department of Zoology, 
Field Museum of Natural History. 1400 South Lake 
Shore Drive. Chicago. IL 60605-2496, U.S.A. 



This review of M. sylvanus is based on study of 
103 specimens (Appendix I) and review of 
relevant literature. ArcView GIS was used to 
prepare maps, and Microsoft Excel was used to 
calculate summary statistics. Reported cranial 
measurements (Table 4) are defined as follows 
(cf. Fooden, 1969, p. 40): Greatest length of 
skull — Most anterior point on rostrum to most 
posterior point on skull. Postrostral length — 
Most inferior point on either orbital margin to 
most posterior point on skull. Rostral length — 
Most anterior point on rostrum to most inferior 
point on either orbital margin. Zygomatic 
breadth — Distance between most lateral points 
on zygomatic arches. 

Names of institutions in which cited specimens 
are preserved are abbreviated as indicated below; 
the number of M. sylvanus specimens studied at 
each institution is indicated by a parenthetical 
notation: 

AMNH American Museum of Natural His- 

tory, New York (7) 



FIELDIANA: ZOOLOGY, N.S., NO. 113, OCTOBER 26, 2007, PP. 1 58 



BM(NH) 


The Natural History Museum. 




London (8) 


CMNH 


Carnegie Museum of Natural His- 




tory. Pittsburgh ( 1 ) 


FMNH 


Field Museum of Natural History, 




Chicago (4) 


IRSN 


Institut Royal des Sciences Natur- 




elles de Belgique. Brussels (17) 


ISR 


Institut Scientifique. Rabat (7) 


MCZ 


Museum of Comparative Zoology, 




Harvard University. Cambridge. 




Massachusetts (2) 


MNHN 


Museum National d'Histoire Nat- 




urelle, Paris (10) 


NHMBa 


Naturhistorisches Museum, Basel 




(1) 


NHMBe 


Naturhistorisches Museum, Bern 




(1) 


NMS 


Natur-Museum Senckenberg, 




Frankfurt (4) 


RMNH 


Nationaal Naturhistorisch Muse- 




um, Leiden (10) 


USNM 


National Museum of Natural His- 




tory, Washington, D.C. (19) 


ZMB 


Museum fiir Naturkunde, Berlin, 




(3) 


ZMUZ 


Zoologisches Museum der Univer- 




sitat Zurich (4) 


ZSBS 


Zoologisches Staatsammlung, Mu- 




nich (5) 



Geographic Distribution 

Extant natural populations of M. syhcmus are 
disjunctly distributed in the Atlas and Rif 
Mountain ranges in Morocco and Algeria 
(Fig. 1); latitudinally the geographic range ex- 
tends from ca. 31'15'N to 36 45'N, and longitu- 
dinally it extends from ca. 7 45'W to 5 35'E. 
Moroccan and Algerian populations are separat- 
ed by a gap that is ca. 700 km wide, and narrower 
gaps separate disjunct populations within these 
two countries. Holocene extinction records 
(Fig. 1, 28 populations) indicate that distribu- 
tional gaps were narrower within historic times 
and further indicate that the range of the species 
recently extended east of Algeria to ca. 10 E in 
neighboring Tunisia; an early reference to M. 
sylvanus in Tunisia has been provided by Her- 
odotus (ca. 484-425 B.C.E.) (McDermott, 1938, 
p. 56). The known elevational distribution of M. 
sylvanus is 400-2300 m (Table 1); half of the 62 



available elevational records are at or above 
18(X) m. No other nonhuman primate is known 
to have ever been sympalric with A/, sylvanus. 

The population of M. sylvanus that currently 
inhabits Gibraltar apparently is not native. 
There is no mention of this species in ancient 
commentaries on Gibraltar (Joleaud, 1931a. 
p. 138; McDermott. 1938. p. 56). Instead, the 
oldest known reference to monkeys on Gibraltar 
is in an 18th century document preserved in the 
British Museum (Sayre. 1862, p. 454; cf. Buffon, 
1789, p. 36); this document indicates that "a 
great quantity" of Barbary macaques were 
introduced into the British garrison in 1740 (cf. 
Busk. 1877, p. 129; Trouessart, 1905, p. 359; 
Morris & Morris, 1966, p. 15; Fa, 1981, p. 74). 
Additional introductions occurred in 1813 and 
1860 (Fa & Lind, 1996, p. 235), but the 
population in 1925 was reduced to three indi- 
viduals (Kenyon, 1938, p. 115), and in 1938- 
1945 major restocking occurred (Fa, 1984b, 
p. 266). A recent study of mtDNA variation in 
the existing Gibraltar population indicated that 
this population includes individuals of both 
Moroccan and Algerian ancestry (Modolo et 
al., 2005, p. 7397). 

Lydekker (1893-1894, p. 118) and Forbes 
(1894, p. 5) indicated that during the 19th 
century, M. sylvanus inhabited southern Spain 
in the vicinity of Gibraltar. This appears to have 
been an error (cf. Kelaart in Newman, 1846, 
p. 1292; Joleaud, 1931a, p. 138), and, in a sub- 
sequent publication, Lydekker (1916, p. 7) omit- 
ted Spain from the species distribution. 

An introduced population of M. sylvanus was 
recently established in Morocco in 1985, when 
more than 40 formerly captive individuals were 
deliberately released near Nador, a northern 
coastal town at ca. 35 OO'N, 3 OO'W (Aulagnier 
& Thevenot, 1986, p. 56; cf. Chivers, 1984, p. vii; 
Anonymous, 1988, p. 50). 



Total Population Estimates 

In 1984, Fa et al. (1984, pp. 86, 91, 99, 101) 
estimated the total extant natural population of 
M. sylvanus to be 14,000-23,000 individuals 
(9,000-17,000 in Morocco, 5,000-6,000 in Al- 
geria). In 1992, Lilly and Mehlman (1993, p. 327; 
cf. Mehlman & Lilly, 1993, p. 334) indicated that 
the population had declined to 10,000-16,000, 
and, in 2005, Modolo et al. (2005, p. 7392; cf. 



FIELDIANA: ZOOLOGY 




Fig. 1. Map of northwestern Africa, showing extant and Holocene locahties of Macaca syhanus; for 
documentation, see Gazetteer, Appendix II. Key to Locality Numbers: MOROCCO and U.K. 1. Goundafa. 2. 
Ourika. Jebel; Ourika, Gued; Tourchte; ZacHa Si Fatma; Zacha Si Fatma, S of. 3. Ouzoud, Cascades d\ 4. Ait 
Mehammed, S of. 5. Tillouguit N"Ait Isha, S of. 6. Ouaouizahrt. MF de. 7. Ighrem. Jebel. 8. Asker, Zaouia; Idrous, 
Jebel; Kousser. Massif du; Tsemair. 9. Ait Sokhmane forest; Azrar; Boutferda vicinity; Haounet, Foret d'El; 
Ijbertene. 10. El Kisba vicinity; Komuch. 11. Asserdoun. 12. Khenifra area. 13. Ajdir, Plateau d'; Ali ou Zrou, 
Jebel; Annocer; Arhbalou N'Irkraouen; Azigza, Aguelmane; Bou Cedre; Bou Igaouerh, Jebel; Imi Ourarn; 
Kerrouchene; Khenifra, 22 km E; Mimejad; Senoual region; Tafechna; Tiguerouguine. 14. Abid, Gued el, gorge; 
Arhbala. N of; Tizi M'Isly, N of. 15. SidiYahya ou Youssef, 8^10 km W of; SidiYahya ou Youssef, S and W of. 16. 
Tounfit, 15-20 km SSE of. 17. Tounfit. N and W of. 18. Tagoulelt vicinity. 19. Itzer, 10-15 km E of; Itzer. W of. 
20. Assaka N"Guam; Zad region. 21. Seheb. 22. Afennourir; Ain Kahla; Ain Leuh; Azrou, 18 km S; El Hammam; 
Guiouane; Gum Er Rbia, Gued. source; Sidi Mguid vicinity; Zaouia de Ifrane. 23. Azrou; Azrou, 5 km S; Azrou, 
above; Azrou Forest; Azrou region; Bou Jirirh; Cedre Gouraud; Ifrane; Michlifene; Ras El Ma. 24. Imouzzer du 
Kandar. vicinity. 25. Taffert; Tahar Souk, N of. 26. Bou Iblane, Jebel; Tamjilt, below. 27. Tahafourt. 28. Rhiata 
mountains. 29. U.K.: Gibraltar; Gibraltar, N end of mountain. 30. Moussa, Jebel. 31. Fath Lemhar. 32. Ahfa- 
Timesh; Buzeitune. Jebel; Derti-Na, N of; El Haouz region; Kelti, Jebel; Sidichmim; Tetouan vicinity; Zarka 
cascade vicinity. 33. Bouhassim, Jebel. 34. Adelma, Gued; Beni Mohammed; Chechaouene, Jebel; El Kelaa; Farda, 
Gued; Kaiat, Jebel; Lakraa, Jebel; Magoo, Jebel; Sidi Salah, Jebel; Sidi Siah; Talaat, Ardhousse; Talassemtane; 
Tasaot. Jebel; Tilljida, Gued; Tisouka, Jebel. 35. Tisirene, Jebel. 36. Targa vicinity. 37. Besene, Bab. 38. Ketama 
vicinity. 39. Tidirhine, Jebel. 40. Tizi Ifri. 41. Al Hoceima. 42. Beni Touzine vicinity. 43. Nador vicinity. 
ALGERIA. 44. Rond-Point des Cedres. 45. Tighret forestry post. 46. Chiffa, Gorges de la. 47. Chiffa, Gorges de la, 
ca. 15 km S of Blida. 48. Chrea. 49. Bou Zegza. Djebel. 50. Palestro, Gorges de; Tala Kitane. 51. Ait Gubane, 
Foret d"; Cheminot; Djurdjura, Djebel; Guessig, Mount; Hotel; Icetcifene; Ras Timedouine; Source des Singes; 
Tala Gulief; Tala Rana; Tigounatine; Tikjda. 52. Akfadou, Foret de; Beni Rhobri, Foret des; Taourirt Irhil, Foret 
de. 53. Bouak. Cap-Cap Carbon, between; Pic des Singes National Park. 54. Akra, Chabet El; Borj Mira; 
Darguinah; Kherrata, Gorges de; Takoucht, Djebel. 55. Babor, Djebel. 56. Adendoun, Djebel; Dar El Gued; 
Guerrouch. Foret Domaniale de; Ziama Mansouria, E of. 57. Kebir, Gued El, gorge. 58. Constantine. 59. Collo, 
Kabylie de. 60. Stora, "Gran"; Stora vicinity. 61. Azzaba, W of. 62. Edough. Djebel. TUNISIA. 63. Tabarqah. 
64. Zaghwan. 



Camperio Ciani & Palentini. 2003, p. 63) re- 
ported a further decline to 10,000. The main 
factor responsible for population decline appar- 
ently is anthropogenic destruction of habitat 
(Fa, 1986a, p. 250; Camperio Ciani et al., 
2005. p. 640; Waters et al., 2007. p. 107). M. 
syhanus has been designated as Vulnerable on 
the lUCN Red List of Threatened Species 
(lUCN, 2006). 



Pelage (Fig. 2) 

In M. sylvanus adults and subadults, prime 
pelage coloration on the dorsal surface of the 
trunk typically is strongly variegated (Fig. 2C), 
pale (pale buffy to golden brown to burnt 
orange) and dark (blackish). This variegation is 
a result of conspicuous banding of individual 
dorsal hairs: in one adult male (usnm 255979, 



FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 



Table 1. Frequency distribution of elevation 
records of Macaca svlvanii.s (see Gazetteer. Appendix 
ID.' 



Elevation (m) 



Number of records 



400 599 


600 799 


800 999 


100{) 


1199 


1200 


1399 


1400- 


1599 


1600 


1799 


1800 


1999 


2000 


2199 


2200-2300 


Total 





2 
1 
7 
9 
4 
6 

21 
9 
1 

62 



' Where elevations were originally reported as 
a range of values, the midpoint of these values is 
tabulated here. 

Gibraltar, zoo specimen), interscapular hair 
length is ca. 80 mm. the basal 50 percent of each 
hair is dark brown, the middle 30 percent is pale 
bufty, and the distal 20 percent is blackish; in 
other adults and subadults. the pale intermediate 
band often is more brightly colored. Premolt 
pelage is faded and drab (Fig. 2A). 

The pelage in one exceptional adult or sub- 
adult female (usnm 258206, Gibraltar, zoo 
specimen) is strikingly erythristic — reddish 
brown on crown, trunk, and appendages. Mor- 
occan specimens that apparently are similarly 
erythristic have been reported by Panouse (1957, 
p. 25: "franchement au roux") and Aulagnier 
and Thevenot (1986, p. 56: "brun-roux sur le 
dos"). According to Bede (1927. p. 28), speci- 
mens obtained in Morocco were generally more 
erythristic than those obtained in Algeria, but 
the purported geographic difference remains to 
be adequately documented (Didier & Rode, 
1936, p. 13; Rode. 1937, p. 158). 

Pelage on the crown is variegated like that on 
the back, but crown hairs tend to be more 
brightly colored. The sides of the face are 
bounded by streaks of dark brown hair, whereas 
the face itself is thinly haired and lightly 
pigmented. The lateral facial crests in 13 speci- 
mens examined are restricted to the posterior 
mandibular region (infrazygomatic crests; Foo- 
den. 1995. p. 19). On outer surfaces of the 
pectoral and pelvic appendages, variegation 
tends to decrease from proximal to distal 
segments, generally becoming pale brown to 
medium brown on dorsal surfaces of the hands 
and feet. The ventral surfaces of the trunk and 



appendages are thinl\ haired, pale buffy to 
whitish. 

At birth, dorsal pelage is blackish, and ventral 
pelage, which is thin, is slightly paler (Fa. 1984c, 
p. 338; cf Carpenlier, 1931, p. 275; Deag & 
Crook, 1971. p. 188; MacRoberts & MacRo- 
berts. 1971. p. 39). The gradual transition from 
blackish infantile pelage to \ariegatcd brownish 
ju\enile pelage begins about age 45 days and is 
completed by about age 145 days. Pelage length 
in juveniles is less than in subadults and adults, 
but the coloration is generally similar. 

Molting occurs during spring and summer and 
usually is complete by September or October 
(Fig. 2C; cf MacRoberts & MacRoberts, 1971, 
p. 39; Merz, 1976, p. 59; Deag, 1984, p. 124). In 
females with new infants, molting usually begins 
about two months after parturition (Fa, 1984c, 
p. 339). The bright new fur appears first on the 
head, arms and forearms; subsequently, it 
appears more caudally and ventrally. 



External Measurements and Proportions 

Head and body length measurements are 
available for a small sample of wild-collected 
M. syhanus specimens, all of which originated 
near Azrou, Moyen Atlas, Morocco (Table 2). In 
this sample, mean head and body length is 
556.8 mm in four adult females and 634.3 mm in 
three adult males; the female/male dimorphism 
ratio is 87.8%. Sexual dimorphism of head and 
body length is similar in six captive specimens or 
specimens of unknown history — 573.0 mm in 
four adult females (amnh 19014, bm[nhi 
11.11.11.1, USNM 258206, 502460) and 630.5 
mm in two adult males (Cabrera, 1932, p. 201; 
Mcz 15296); in these specimens, the female/male 
length ratio is 90.9%. No information is available 
concerning geographic variation of head and 
body length in M. sylvanus. 

Mean body weight in wild-collected adults is 
9.9 kg in 18 females and 14.5 kg in 33 males 
(Table 2; cf Roberts, 1978, p. 233; Fa, 1984c, 
pp. 341 346; 1989, p. 58); the female/male di- 
morphism ratio for this volumetric measurement 
is 68.3%. Mean weight in four adult females 
collected in Morocco (9.2 ± 0.82 kg) is not 
significantly different {P > 0.05, /-test) from that 
in 14 females probably trapped in Algeria (10.1 
± 1 .03 kg); weights of Moroccan adult males are 
unavailable (cf Table 2, footnote 4). 



FIELDIANA: ZOOLOGY 




Fig. 2. Adult pelage in Mucaca \yl\aiiiis. A. Male, prcmoll. B. Male, molting (note new pelage on head and 
shoulders). C. Estrous female, postmolt. Photographed 29 September 2006 at Cedre Gouraud, Morocco, by G. 
Fooden; image in figure B is photographically reversed. 

Relative to adult size in M. sylvanits, ears in foot in infants is only ca. 60% of that in 

infants are larger than hind feet (Table 2). adults. 

Length of the ear in infants is ca. 80% of An inconspicuous vestigial tail is variably 

that in adults, whereas length of the hind present in M. sylvcmus (Fig. 3; Table 3; cf 



FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 



Table 2. External measurements and proportions in wild-collected Macacu syhcinus. 











Relative hind 




Relative 




Head and b(>d> 


Weight 


Mind foot 


foot length 


Kar length 


ear length 


.\ge/se\ class 


length (mm) 


(kg) 


length (mm) 


(HI/MB KM)) 


(mm) 


(K/HB X 100) 


Infants 


266.5 


1.0 


101.5 


38.2 


40.0 


15.0 




246 287 


0.9 1.0 


96-107 


37.3 39.0 


36 44 


14.6 15.3 




(2) 


(2) 


(2j 


(2) 


(2) 


(2) 


Juveniles 


521.0 ± 28.16 


7.1 ± 1.31 


159.0 ± 13.89 


30.5 ± 1.10 


45.0 ± 6.24 


8.6 ± 1.00 




500 553 


6.2-8.6 


150 175 


29.4-31.6 


38 50 


7.5-9.4 




U) 


(i) 


U) 


(i) 


(i) 


(i) 


Subadult males 


604.0 














559 649 




182 


28.0 


51 


7.9 




(2) 


— 


(/) 


(/) 


(/) 


(/) 


Adult females 


556.8 ± 11.47 


9.9 ± 1.04 


167.8 ± 1.71 


30.1 ± 0.68 


48.8 ± 1.26 


8.8 ± 0.10 




540-566 


8.0-12.0 


166 170 


29.3 30.9 


47 50 


8.7-8.9 




(4) 


{/<V) 


(4) 


(4) 


(4) 


i4) 


Adult males 


634.3 ± 25.72 


14.5 ± 1.75 


179.0 ± 7.94 


28.3 ± 1.85 


51.3 ± 1.15 


8.1 ± 0.42 




605-653 


10.0 18.0 


170-185 


26.4-30. 1 


50 52 


7.8 8.6 




(3) 


U-^f 


(i) 


(3) 


(3) 


(3) 



' Mean ± SD (where n > 2), extremes, and sample size (italicized figures in parentheses). 

" Most measurements are of specimens collected near Azrou. Morocco (bm(NH). usnm). Exceptions are weights of 
14 adult females and 33 adult males that were reported by Delson et al. (2000, pp. 144-145); these 47 adults 
probably were trapped and weighed in Algeria (cf. Scheffrahn et al., 1993. p. 383; 1996. abstract no. 240). 
Measurements of isnm specimens are from specimen tags and collectors' field catalogs. 

^ Dental specifications: infants, deciduous teeth only; juveniles, some permanent teeth erupted; subadults. M3 in 
females or C in males incompletely erupted; adults, all permanent teeth completely erupted. 

■* Excludes USNM 476781 (juvenile) and usnm 476791 (weight recorded as ">10 kg") (cf. Delson et al., 2000. 
p. 144). 



Lydekker. 1893-1894. pp. 117 119); Aristotle 
(d"A. W. Thompson translation, 1984, p. 799) 
accurately described the tail vestige in this species 
as "... a very small one — a sort of hint of a tail." 
In a series of 13 Moroccan specimens that were 
carefully examined by the collectors, a tail 
vestige, varying in length from 4 mm to 
22 mm, was detected in eight individuals (62%); 
although the sample size is small, the data 
suggest that the tail vestige might be more 
frequent in males (86%) than in females (33%). 
Tail vestiges of similar length are known to occur 
in three captive males (Hill, 1974, p. 622, 
18.7 mm: fmnh 60716, 9 mm [Fig. 3]; mcz 
15206, 22 mm). Relative tail length probably is 
greater in infants than in postinfants (Table 3), 
which suggests that the postnatal growth rate of 
the tail is less than the postnatal growth rale of 
the head and body. 



Caudal Vertebrae 

Caudal vertebrae also are vestigial in M. 
sylvamis (Fig. 4), and, as previously noted by 



Waldeyer (1896, p. 783) and Schultz (1925, 
pp. 249-250), these vertebrae do not extend into 
the external tail; for this reason, Waldeyer and 
Schultz have characterized the tail vestige in M. 
sylvamis as "soft." The number of caudal 
vertebrae in this species varies from two to five 
(Appendix III), with a modal number of three 
(13 of 29 specimens reported). Reduction in 
number of caudal vertebrae in M. sylvanus 
apparently exceeds that in humans, where the 
modal number of caudal vertebrae is four 
(Schultz & Straus, 1945. p. 613). 



Cranial Characters 

Mean greatest length of skull in wild-collected 
M. sylvanus is 118.7 mm in four adult females 
and 135.5 mm in five adult males (Figs. 5, 6; 
Table 4); the female/male dimorphism ratio is 
0.88. Available evidence does not permit analysis 
of geographic variation of skull size in this 
species (cf. Table 4, footnote 4). 

In wild-collected M. sylvanus, mean relative 
zygomatic breadth increases from 60. 9%^ in 



FIELDIANA: ZOOLOGY 



10mm 





Fig. 3. Tail vestige (indicated by arrow) in Macaca sylvatius (fmnh 60716, infant male, zoo specimen); tail 
length is 9 mm. (Photograph by R. Banasiak.) 



infants to 69.2% in adult males, and mean 
rostral-postrostral ratio increases from 23.8% 
in infants to 59.0% in adult males (Table 4). As 
in other macaques, postnatal relative growth of 
the rostrum greatly exceeds that of the zygomatic 
arches. 



A sagittal crest is relatively rare in M. syhcinus. 
In adult male skulls examined, this crest is absent 
in four wild-collected specimens and present in 
only two (Mcz 15296, mnhn CG 1831/835) of 
nine captive specimens or specimens of unknown 
history (overall frequency, 2/13 = 15.4%). 



FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 



Table 3. Tail length and relative tail length in Macaca sylvanus specimens collected near Azrou, Mo\en Atlas, 
Morocco; measurements from specimen tags and collectors' field catalogs. 









Head and body 


I ail length" 


Relatiu- tail length 


ISNM No. 


Age' 


Sex 


length (mm) 


(mm) 


(I/HB ■ UK)) 


476788 


Infant 


Male 


246 


15 


6.1 


476789 


Infant 


Male 


287 


15 


5.2 


476790 


Juvenile 


Female 


510 





— 


478329 


Ju\cniie 


Female 


500 


15 


3.0 


476781 


Juvenile 


Male 


553 


15 


2.7 


476784 


Subaduit 


Male 


649 


22 


3.4 


476782 


Adult 


Female 


561 





— 


476783 


Adult 


Female 


566 


4 


0.7 


476786 


Adult 


Female 


540 





— 


476787 


Adult 


Female 


560 





— 


476780 


Adult 


Male 


605 


20 


3.3 


476785 


Adult 


Male 


653 


22 


3.4 


476791 


Adult 


Male 


645 





— 



' For dental specifications, see Table 2. 

- Cf. MacRoberls and MacRoberts. 1971. p. 39; Burton and de Pelham. 1979, p. 109 (apparent decimal point 
placement errors in this publication; tail length reported as "0.5 mm" and "0.53 mm" in two infant specimens). 



Based on analyses of 27 cranial dimensions. 
Pan and Oxnard (2004, p. 367) have reported 
that cranial differences between M. sylvanus (21 
adult specimens) and 10 Asian macaque species 
(303 adult specimens) greatly exceed correspond- 
ing differences among the 10 Asian species. 
Unfortunately, the provenance of ihe M. sylva- 
nus specimens (and non-M. sylvanus specimens) 
studied in this research project is not specified 
(cf. Table 4); some artificially provisioned indi- 
viduals might have been included (cf. Fooden & 
Aimi. 2005. p. 17). 



Molecular Biology and Genetics 
DNA Variation 

Virtually all studies of interspecific DNA 
variation within the genus Macaca have in- 
dicated that M. sylvanus is the sister group of 
Asian macaque species (Hayasaka et al., 1996. 
p. 1050: Morales & Melnick, 1998. pp. 15-16; 
Deinard & Smith. 2001. p. 50; Tosi et al., 2003, 
pp. 1427, 1430; Modolo et al., 2005. p. 7394; Liu 
& Pearl. 2007, p. 411; Ziegler et al., 2007, 
p. 811). A recent exception is Tosi et al.'s 
(2003, p. 1426) study of Y-chromosomal DNA, 
in which the phylogenetic relationship of M. 
sylvanus was unresolved. 

The most comprehensive survey of intraspe- 
cific DNA variation in M. sylvanus has been 



published by Modolo et al. (2005, p. 7393). In 
this study, a 468-bp sequence of hypervariable 
region 1 of the mtDNA control region was 
analyzed in 68 samples of Gibraltar monkeys 
(artificially introduced population; see above. 
Geographic Distribution), 116 samples of Mor- 
occan monkeys (3 localities), and 96 samples of 
Algerian monkeys (4 localities) (Table 5). Of the 
24 haplotypes detected in non-Gibraltar samples, 
nos. 1-15 were restricted to Morocco and 
nos. 16-24 were virtually restricted to Algeria 
(cf. Pastorini et al.. 2000, p. 166; Crouau-Roy et 
al., 2001, p. 157; von Segesser, 2002. p. 161); the 
sole exception is haplotype no. 16, which was 
detected in 25 Akfadou (Algeria) samples and 
four Rif (Morocco) samples. Modolo et al. 
(2005, pp. 7394, 7397) plausibly concluded that 
the anomalous geographic distribution of hap- 
lotype no. 16 probably is a result of Algerian 
monkeys having been artificially transported and 
released in the Rif mountains. Haplotype evi- 
dence (Table 5) also indicated that the Gibraltar 
population has resulted from artificial introduc- 
tion of monkeys from the Rif (Morocco) and 
Akfadou (Algeria) regions (Modolo et al., 2005, 
p. 7397). 

Phylogenetic analysis indicates that Algerian 
mtDNA haplotypes are more divergent than 
Moroccan haplotypes (Fig. 7); in fact, Algerian 
haplotypes at Pic des Singes (haplotype no. 18) 
and Kherrata (haplotypes nos. 19 22) are more 
similar to Moroccan haplotypes than they are to 
other Algerian haplotypes (cf. Scheffrahn et al.. 



FIELDIANA: ZOOLOGY 




Fig. 4. Ventral view of sacrum and caudal vertebrae (Cd 1-Cd 3) in Macaca sylvanus (fmnh 47398, adult male, 
zoo specimen); vestigial Cd 3 is asymmetrically fused to Cd 2. (Photograph by J. Weinstein.) 



1996, abstract no. 240). Modolo et al. (2005, 
p. 7396) have indicated that the present disjunct 
clustering of mtDNA haplotypes could be 
a product of Pleistocene and Holocene climate 



changes that resulted in periodic forest fragmen- 
tations and interconnections. Based on fossil 
evidence and the date of the Messinian Salinity 
Crisis (Delson, 1980a. p. 16; Tosi et al., 2003, 



FOODEN; SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 




Fig. 5. Skull of adult female Macucu sylvanm: usnm 476782. Azrou. 1 8 km S of, Morocco. (Photographs by E. 
M. Santana.) 

p. 1424), Modolo el al. (2005, p. 7393) have differentiated within the past 1.85 million years 

postulated that M. syhanus originated ca. 5.5 (Table 6). 

million years ago (Ma) and have inferred that Microsatellite variation within and between 

extant clades and subclades of M. sylvanus four isolated Algerian populations of M sylva- 



10 



FIELDIANA: ZOOLOGY 




Fig. 6. Skull of adult male Macaca sylvamis: usnm 476785, Azrou, 18 km S of, Morocco. (Photographs by E. 
M. Santana.) 



FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 



11 



Table 4. Cranial measurements'' and proportions in wild-collected Macuca sylvanus; except for one adult 
male obtained at Gorges dc la ChifTa. Algeria, all specimens were collected near Azrou. Morocco (see footnotes 4 
and 5). 





(Jreatest length 


Relative zygomatic 


Postrostral 


Rostral-postrostral ratio 


Age/sex class"* 


of skull (mm) 


breadth (ZB/GL x 


100) 


length (mm) 


(R/PR X 100) 


Infants 


80.3 


60.9 




71.1 


23.8 




77.6 -83.0 


60.4 61.4 




68.4 73.7 


23.3-24.4 




(2) 


(2) 




(2) 


(2) 


Juveniles 


113.0 


66.6 




82.6 


48.0 




109.8-116.3 


65.7-67.5 




81.2-83.9 


43.7-52.2 




(2) 


<-^ . 




(2) 


(2) 


Adult females 


118.7 ± 5.48 


69.4 ± 2.05'* 




85.9 ± 1.66 


49.6 ±3.12 




113.7-125.7 


68.2-72.4 




84.1-87.5 


47.4-54.2 




i4) 


(4) 




(4) 


(4) 


Subadult males 


124.3 


67.6 




90.7 


56.5 




118.2-130.5 


66.0-69.2 
( '') 




(/) 


(/) 


Adult males 


135.5 ± 2.27^ 


69.2 ± 1.00-' 




91.7 ± 0.95 


59.0 ± 2.47 




132.7-137.7 


68.0-70.5 




90.5-92.7 


55.6 61.6 




(5) 


{5) 




(-/) 


(•/) 



' For definition of measurements, sec above. Materials and Methods. 

" Mean ± SD (where sample size exceeds two), extremes, and sample size (italicized figures in parentheses). 

' Dental specifications: infants, deciduous teeth only; juveniles, some permanent teeth erupted: subadults, M3 in 
females or C in males incompletely erupted: adults, all permanent teeth completely erupted. 

"* 135.3 ± 2.58 mm in four adult males collected in Morocco; 136.2 mm in one adult male collected in Algeria 
(measurement of Algerian specimen provided by D. Brandon-Jones). 

^ Zygomatic breadth of Algerian specimen estimated by D. Brandon-Jones. 



nus was studied by von Segesser et al. (1999, 
p. 436). In a sample of 159 monkeys, six of the 
seven loci tested were found to be polymorphic, 
with 4-12 alleles per locus. Populations and 
population subgroups were genetically differen- 
tiated with respect to these loci. Genetic distance 
was significantly correlated with geographic 
distance within a population — but not between 
populations. Crouau-Roy et al. (2001, p. 157), 
who also studied microsatellite variation in M. 



sylvanus, found 12 alleles specific to Moroccan 
samples and 21 alleles specific to Algerian 
samples. 

Karyology 

The chromosomes of M. sylvanus have been 
studied — with partly divergent results — by Stan- 
yon et al. (1980, p. 150), Dutrillaux et al. (1982, 
p. 97), Tihy et al. (1996, p. 10), and Morescalchi 



Table 5. Mitochondrial DNA haplotype frequencies in Gibraltar and North African samples of Macacci 
sylvanus (Modolo et al.. 2005. p. 7393). 



Localitv 



Haplotype code numbers 



A' 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 









U.K. 




Gibraltar 


68 


24 


8 
Morocco 


36 


Rif 


28 


4 


20 


4 


Moyen Atlas 


69 4 


16 3 1 


12 10 1 16 2 3 


1 


Haut Atlas 


19 


6 6 3 


4 

Algeria 




Akfadou 


25 






25 


Djurdjura 


54 








Pic des Singes 


12 








Kherrata 


5 









45 9 



7 5 



2 1 I I 



12 



FIELDIANA: ZOOLOGY 



Rif 

Moyen Atlas 
Haut Atlas 
Kherrata 



Mor. 
Mor. 
Mor. 
Alg. 



— Pic des Singes (hap. no. 18) Alg. 

- Djurdjura Alg. 

- Akfadou Alg. 



- Pic des Singes (hap. no. 17) Alg. 

Fig. 7. Phylogenetic relationships among Moroc- 
can (Mor.) and Algerian (Alg.) populations of Macaca 
syhamis, as indicated by mtDNA haplotypes (maxi- 
mum likelihood tree, condensed from Modolo et al., 
2005. p. 7394); the Pic des Singes sample includes two 
haplotype subgroups. 

et al. (1998, p. 102). Stanyon et al. (1980, p. 150) 
found that chromosomes in M. sylvanus are 
generally similar to those in M. nnilatta, except 
that banding patterns in four chromosomes 
(nos. 1, 4, 9, and 16) are polymorphic in M. 
sylvanus but not in M. mulatta. Dutrillaux et al. 
(1982, p. 107) reported that chromosomes in M. 
sylvanus are similar to those in other macaques 
(except M. fasclcularis) and also are similar to 
those in species of Papio and Lophocehus. Tihy et 
al. (1996, p. 10) reported that chromosome 
no. 18 in M. sylvanus is homologous to chromo- 
some no. 13 in humans; a centromeric shift 
probably occurred during the course of this 
chromosomal transition. And Morescalchi et al. 



(1998, p. 103) reported that chromosome no. 2 
in M. sylvanus has a clear band adjacent to the 
centromere that does not occur in other species 
of Macaca but does occur in species of Papio; 
these authors also found that human chromo- 
some probes nos. 7 and 21 hybridized to M. 
sylvanus chromosome no. 2 (cf. Wienberg et al., 
1990, p. 347). 

As in other members of the tribe Papionini, 
the diploid chromosome number in M. sylvanus 
is 42. 



Natural History 

Habitats 

Habitats occupied by M. sylvanus include 
cedar, fir, and oak forests (evergreen oak and 
deciduous oak), scrub, grasslands, and rocky 
ridges dominated by herbaceous vegetation 
(Table 7); although most extant M. sylvanus 
populations inhabit cedar forests, this might 
reflect current habitat availability rather than 
habitat preference (Fa, 1984e, pp.281, 284). 
Climate in the range of this species is Mediter- 
ranean, with warm dry summers and relatively 
wet winters (Trewartha, 1978, p. 8); at elevations 
above 1500 m — i.e., well within the elevational 
range of M. sylvanus (Table 1) — snow falls in 
winter, and snow accumulations can persist on 
mountain summits until mid-May (Fa, 1982, 
p. 50; Mehlman, 1984, p. 172). Alvarez and 
Hiraldo (1975, p. 256) have reported that mon- 
key populations in the Rif mountains migrate to 
lower elevations during winter. 



Table 6. Estimated dates of divergence of Macaca svlvamts sister groups, based on mtDNA haplotype 
relationships (Modolo, 2005, p. 7394, table 4). 



Sister groups (see Fig. 7) 



Estimated divergence dates (Ma)' 



Morocco/Kherrata/Pic des Singes (hapl. Djurdjura/Akfadou/Pic des Singes 1.24—1.85 

no. 18) clade (Morocco + Algeria) (hapl. no. 17) clade (Algeria) 

Morocco clade Kherrata/Pic des Singes (hapl. no. 18) 1.22-1.62 

clade (Algeria) 

Djurdjura clade (Algeria) Akfadou/Pic des Singes (hapl. no. 17) 0.78-1.01 

clade (Algeria) 

Kherrata subclades (Algeria) 0.064-0.359 

Morocco subclades 0.110-0.282 

Akfadou subclades (Algeria) 0.070-0.078 

Djurdjura subclades (Algeria) 0.070-0.078 



Million years ago. 



FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 



13 



Table 7. Habitats of Maiacu svlvamts. 



Geographic area 



Dominant vegetation 



Scientific name 



References' 



Morocco 
Haiit Atlas 
Moyen Atlas 



Rif 



Algeria 
ChifTa 

Tigounatine 
Icetcifere 
Akfadou 
Kerrata 



Scrub 


— 


/, 2 


Atlantic cedar 


Cedrus atlaniica 


1. 3. 4 


Holm oak" 


Quenus ilex 




Mixed cedar oak 


— 




Grassland 


— 




Spanish fir 


Abies pinsapo 


2. 5. 6. 


Atlantic cedar 


Cedrus atlantica 




Holm oak' 


Qiiercus ile.x 




Mixed oak 


— 




Scrub 


— 




Holm oak" 


Quenus ilex 


6 


Mixed oak 


— 




Mixed cedar-oak" 


— 


9. 10 


Herbaceous 


— 


9, 10 


Mixed oak' 


— 


10. 11 


Holm oak" 


Quenus ilex 


6 


Mixed oak 


— 





' Kev to reference numbers: 1. Deag and Crook, 1971. p. 186. 2. Taub, 1977. p. 110. 3. Menard, 2002, p. 95. 4. 
Camperio Ciani et al.. 2005, p. 639. 5. Alvarez and Hiraldo. 1975, pp. 254-257. 6. Fa et al., 1984. pp. 86. 100. 105. 
7. Mehlman, 1984, p. 165. 8. Fa, 1986a, p. 222. 9. Menard and Vallet, 1986, pp. 173, 174. 10. Menard, 2002, p. 95. 
//. Menard. 1985, p. 452; 2002, p. 95. 

" Evergreen. 

"^ Deciduous. 



Terrestriality/.\rboreality 

Macaca sylvanus spends most of the daylight 
hours on the ground (Table 8); reported mean 
frequency of daytime terrestriality varied from 
68% to 83% in Morocco and from 58% to nearly 
100% in Algeria. Infants and juveniles tended to 
be less terrestrial than adults (Merz, 1976, p. 60; 
Deag, 1985, p. 408; Menard, 1985, p. 466; 
Menard, & Vallet, 1986, p. 176; Machairas et 
al., 2003, p. 189); in populations studied at 
Afennourir and Cedre Gouraud ( = Gouroud), 
Morocco, juveniles were 70% terrestrial and 
adult males were 81% terrestrial (Machairas et 
al., 2003, p. 189). Frequency of terrestriality also 
varied seasonally (Deag, 1985, pp. 408-409; 
Menard & Vallet, 1986, p. 190; 1997, p. 292). 
This species flees into trees to escape danger 
(Merz, 1976, p. 60; Deag, 1985, p. 408), and it 
sleeps in trees (Taub, 1977, p. 128; de Turkheim 
& Merz, 1984, p. 247; Mehlman, 1989, p. 285; 
Hammerschmidt et al., 1994, p. 279) or in caves 
on rocky cliffs (MacRoberts & MacRoberts, 
1971, p. 45; Alvarez & Hiraldo, 1975, p. 258; 
Fa et al., 1984, p. 94; Mehlman, 1984, pp. 190- 
191). 



Group Size and Composition 

Reported size in 68 natural groups of M. 
sylvanus averaged 27.1 individuals and varied 
from seven to 88 individuals (Appendix IV). 
Limited available evidence has suggested that 
group size might average smallest (18.3 individ- 
uals) in the Moroccan Rif, largest (50.2 individ- 
uals) in the Algerian Grande Kabylie, and 
intermediate (28.6 individuals) in the Moroccan 
Moyen Atlas (Appendix IV). Group size gener- 
ally is fairly stable in this species, but it was 
unstable in an Algerian population that inhabits 
a rocky habitat in the Djurdjura mountains 
(Menard et al., 1985, p. 72; 1990, p. 169; 
Menard, 2002, pp. 96, 98); in this marginal 
habitat, groups frequently split into subgroups 
that subsequently reunite in various combina- 
tions. Solitary males, as far as 7 km distant from 
the nearest group, have been reported in the 
Moroccan Rif (Mehlman, 1985, p. 204; 1986, 
p. 75). Provisioned semi-free-ranging groups, 
which sometimes have included as many as 200 
members (de Turckheim & Merz, 1984, p. 243; 
Kuester & Paul, 1992, p. 194), apparently can 
become larger than natural groups. 



14 



FIELDIANA: ZOOLOGY 



Table 8. Frequency of terrestriality in Maccica sylvamts populations. 









Mean frequency 




Locality 




Sample size 


of terrestriality 


References' 






Morocco: Moyen Atlas 






Ai'n Kahla (cedar) 




11,202 observations 


83% 


I 


Afennourir (cedar) 




317 hr 


68% 


2 


Cedre Gouraud" (cedar-oak) 




267 hr 

Algeria: Grand Kabylie 


77% 


2 


Tigounatine (cedar-oak) 




2,587 scans 


68% 


3 


Akfadou (oak) 




2,608 scans 


58% 


3 


Icetcifere (mountain ridge) 




France: semi-free-ranging 


-100% 


4 


Kintzheim (mixed European 


forest) 


— 


>50% 


5 



' Key to reference numbers: /. Deag, 1985, p. 409. 2. Machairas et al., 2003, p. 189. 3. Menard and Vallet. 1997, 
p. 292; cf. Menard, 1985, p. 466; Menard and Vallet, 1986, p. 176. 4. Menard and Vallet, 1986, p. 190. 5. Merz, 
1976, p. 60. 

" Alternate locality name: Gouroud. 



Within natural groups of M. sylvcmus, the 
number of adult males apparently averages only 
slightly less than the number of adult females 
(Appendix IV). The known ratio of adult males 
to adult females averaged 0.70 in the Moroccan 
Moyen Atlas, 0.99 in the Moroccan Rif, and 
0.84 in the Algerian Grande Kabylie. 

Group Fission 

A M. sylvanus group in the Algerian Grande 
Kabylie that increased in size from 38 members 
in 1983 to 88 members in 1988 permanently 
subdivided in 1989 into three daughter groups 
(50, 24, and 13 members; 1 member disappeared) 
(Menard & Vallet, 1993a, p. 483; 1993b, p. 105; 
Lathuilliere et al., 2004, p. 176). As noted above, 
88 is the maximum group size known in natural 
groups of M. sylvanus, which suggests that 
fission could be inevitable in groups that 
approach this size. 

Fission also has been reported in semi-free- 
ranging European groups of M. sylvanus (Prud'- 
Homme, 1991, p. 10; Fa & Lind, 1996, p. 237; 
Kuester & Paul, 1997, p. 946; Fischer, 1998, 
p. 800; KummerU & Martin, 2005, p. 1232). 
Female matrilines generally remained intact 
during the course of these fission events, which 
usually began during the mating seasons. 

Home Range Area and Population Density 

Available evidence suggests that home range 
area in natural groups of M. sylvanus averages 
smallest (18.4 ha) in the Moroccan Moyen Atlas, 



largest (804.5 ha) in the Moroccan Rif, and 
intermediate (279.7 ha) in the Algerian Grand 
Kabylie (Table 9). Home ranges of adjacent 
groups frequently overlapped — 100% overlap 
reported for some groups — in all three areas 
(Deag & Crook, 1971, p. 188; Rumsey & 
Whiten, 1972, p. 562; Menard et al., 1986, 
p. 37; Mehlman, 1989, p. 282). Of 13 observed 
visual contacts between neighboring groups, only 
three (23%) were agonistic (Mehlman & Parkhill, 
1988, p. 33; cf. Deag & Crook, 1971, p. 189). 
Linear day ranges of >2 km have been reported 
for two Algerian groups (Menard & Vallet, 1997, 
p. 297). 

Population density in M. sylvanus apparently 
attains its maximum in relatively undisturbed 
cedar or mixed cedar/oak habitats, where densi- 
ties of 19-70 individuals per km" have been 
reported (Table 10). As expected, degraded 
habitats, particularly those in the Moroccan 
Rif, support population densities that are much 
lower — 0.37-4.50 individuals per km" (cf. Mehl- 
man, 1989, p. 286; Dell'Amico et al., 2006, 
p. 292). Two semi-natural populations main- 
tained in France have reached densities of 
1,500 and 2,600 individuals per km" (de Turck- 
heim & Merz, 1984, p. 245). 

Diet 

The diet of natural populations of M. sylvanus 
includes a wide variety of gymnosperms and 
angiosperms (Fa, 1984d, pp. 348-355) (Ta- 
ble 11). In the Moroccan Moyen Atlas, 107 
species of food plants have been reported 



FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 



15 



Table 9. Home range area in natural populations of .\finiiiii sylvaniis. 



Sample area 



Group size 



Home range area (ha) 



References 



Ain Kahla 
Bou Jirirh 
Means (weighted) 

Beni Hessane: Bouhassim 
Rhomara: Talssemtane 
Means (weighted) 

Akfadou 

Djurdjura: Mount Guessig 
Djurdjura: Tigounatine 
Means (weighted) 



Morocco: Moven Atlas 

32- (25 39) 15.1- (11.7- 18.6) 

40 25.0 

34.7 18.4 
Morocco: Rif' 

27 1200.0 

28.8-* (12 59) 725.4^(307-901) 

28.5 804.5 
Algeria: Grande Kabylie 

39-45 424 

>37 39 

46 73 376 

>46.2 279.7 



' Kev to references: /. Fa. 1986d. p. 202. 2. Drucker, 1984, p. 139; Fa, 1986d, p. 202. 3. Mehlman, 1989, p. 282. 
4. Menard and Vallet. 1996. pp. 123-124. 5. Menard et al.. 1990. p. 169. 

■ Mean value of two groups; extreme values indicated in parentheses. 

^ Cf. Rumsey and Whiten. 1972. p. 562. 

"* Mean value of five groups; extreme values indicated in parentheses. 



(Drucker. 1984, p. 142; cf. Deag, 1983, p. 573); 
in the Moroccan Rif. 100 species have been 
reported (Mehlman, 1988. pp. 472. 477; cf. Fa. 
1983a. p. 84); and in the Algerian Grand 
Kabylie, 130 species have been reported (Menard 



& Vallet, 1986, pp. 180, 190; 1996, pp. 120, 125; 
cf. Menard, 1985, pp. 460-463). The 100 species 
exploited as food in the Moroccan Rif constitut- 
ed 51 9f of 195 species of seed plants identified as 
present in that area, and, similarly, the 130 



Table 10. Population density in Mcucica sylvaiiii.s 



Sample area 


Dominant vegetation 


Survey year 


Density (individuals/km") 


References' 




Morocco: 


Moyen Atlas 






Afennourir 


Cedar (protected) 


1994^1995 


19.3 


1 


Ain Kahla 


Mixed cedar/oak 


1968-1969 


60-70 


2 


Ain Kahla 


Mixed cedar/oak 


1973-1974 


40 


3 


Azrou vicinity 


Mixed cedar/oak 


1994 


28.19 


4 


Azrou vicinity 


Mixed cedar/oak 
(degraded) 


2002 


7.26 


4 


Bou Jirirh 


Cedar (undamaged) 


1980-1981 


50 


5 


Cedre Gouraud" 


Mixed cedar/oak 
(degraded) 


1994^1995 


7.8 


1 


Oued-el-Abid 


Mixed oak/juniper 


1984 


-11 


6 




Morocco: Rif 






Anjcra: Moussa. Jebel 


Scrub 


1980 


1.09 


7 


Beni Hessane: Bouhassim 


Mixed oak 


1980 


0.37 


7 


Beni Zaid 


Scrub 


1980 


0.60 


7 


Rhomara 


Various 


1980 


1.08-4.50 


7 


Rhomara: Talassemlane 


Fir 

Algeria: G 


1981 1982 
rande Kabylie 


8.3 


8 


Akfadou 


Deciduous oak 


1990 


12.5 


9 


Djurdjura: Tigounatine 


Mixed cedar/oak 


1990 


28.2 


9 



' Key to references: /. Machairas et al., 2003, p. 189; cf. Camperio Ciani and Machairas, 2003, p. 124. 2. Deag. 
1977. p. 269; 1984, p. 123. 3. Taub, 1977, p. 110. 4. Camperio Ciani et al., 2005, p. 638. 5. Drucker, 1984, p. 136; 
Fa. 1986d, p. 202. 6. Fa, 1986b, p. 33. 7. Fa. 1982. p. 62. H. Mehlman, 1984, p. 175. 9. Menard and Vallet, 1996, 
p. 124. 

- Alternate locality name: Gouroud. 



16 



FIELDIANA: ZOOLOGY 



Table 11. Number of gymnosperm and angiosperm (seed-plant) species consumed by Macaca 
sylvciiilis populations. 



Sample area 



Number of seed-plant 
species eaten 



References 



Ai'n Kahla 
Bou Jirirh 


70 
107 


Djebala region 
Lakraa. Jebel 


46 
100' 


Akfadou 
Icetcifere 
Tigounatine 


77 

31 

130- 



Morocco: Moyen Atlas 
Deag. 1983, p. 573 
Drucker. 1984, p. 142 

Morocco: Rif 
Fa, 1983a, p. 84 
Mehlman, 1988, pp. 472, 477 
Algeria: Grande Kabylie 
Menard, 1985, pp. 460-463; Menard & Vallet, 1996, pp. 120. 125 
Menard & Vallet, 1986, p. 190 
Menard & Vallet, 1996, pp. 120, 125 



Total seed-plant species identified as present in this sample area: 195 (51.3% consumed). 
Total seed-plant species identified as present in this sample area: 271 (48.0% consumed). 



species exploited in the Algerian Grand Kabylie 
constituted 48% of 271 species identified as 
present in that area. Barbary macaques also 
consume fungi, lichens, mosses, and animal 
prey (Deag, 1983, p. 571; Fa, 1984d, p. 355; 
Menard, 1985, p. 452). Agricultural crops are 
known to have been raided by Barbary macaques 
at least since the early 16th century (Leo 
Africanus, 1896 edition, p. 948; cf Mehlman, 
1988, p. 477). 

Plant parts that are eaten by M. sylvanus 
include flowers, fruits, seeds, seedlings, leaves, 
buds, bark, gum, stems, roots, bulbs, and corms 
(Fa, 1984d, p. 348; Mehlman, 1988, p. 472; 
Menard & Qarro, 1999, p. 130). Because dietary 
composition varies seasonally (Deag, 1983, 
p. 571; Mehlman, 1984, p. 192; Menard, 1985, 
p. 455; Menard & Vallet, 1986, p. 177), compar- 
ative quantitative evaluation of dietary compo- 
nents requires study periods that extend over at 
least 12 months. Studies of that duration have 
indicated that seeds and leaves are the main 
foods (ca. 60-75%) of Barbary macaques in the 
Algerian Grande Kabylie (Table 12). Other 
studies have suggested that fruits could consti- 
tute a more significant part of the diet in the 
Moroccan Moyen Atlas (Deag, 1983, p. 574; 
Drucker, 1984, p. 142; Menard & Qarro, 1999, 
p. 130). 

Animal prey consumed by M. sylvanus in- 
cludes snails, earthworms, scorpions, spiders, 
centipedes, millipedes, grasshoppers, termites, 
water striders, scale insects, beetles, butterflies, 
moths (including larvae), ants (including nests), 
and tadpoles (Fa, 1983a, p. 84; 1984d, p. 355; 



Mehlman, 1984, p. 193; 1988, p. 476; Menard & 
Vallet, 1986, p. 187). Semi-free-ranging monkeys 
have been observed to pursue and/or catch birds, 
squirrels, and young rabbits, but never to eat 
them (Kaumanns, 1978, p. 59; de Turckheim & 
Merz, 1984, p. 248); mice apparently are ignored. 

Feeding reportedly occupied an average of 
24% and 25% of daytime hours at two localities 
in the Moroccan Moyen Atlas (Machairas et al., 
2003, p. 193; cf Fa, 1986c, p. 216) and also 
occupied 24%. and 25%> of daytime hours at two 
localities in the Algerian Grande Kabylie (Me- 
nard & Vallet, 1997, p. 291). 

In winter, water has been obtained by eating 
snow (de Turckheim & Merz, 1984, p. 249; 
Mehlman, 1988, p. 482). During other times of 
water shortage, M. sylvanus has stripped the 
bark of cedar trees in order to obtain retained 
moisture (Camperio Ciani et al., 2001, p. 263). 

Predators 

Under natural conditions, the leopard proba- 
bly was a major predator of M. sylvanus when 
these two species were broadly sympatric (Au- 
lagnier & Thevenot, 1986, p. 56; cf Fa, 1986b, 
p. 31). Large eagles presumably also were and 
are predators, because Barbary macaques re- 
spond to eagles' approach with alarm calls 
(Mehlman, 1984, p. 191); domestic dogs elicit 
a similar response, but foxes and jackals do not 
(cf Menard et al., 1986, p. 36). 

In semi-free-ranging European populations, 
M. sylvanus gave alarm calls in response to the 
appearance of snakes, raptorial birds, and dogs 



FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 



17 



Table 12. Allocation of feeding time to various dietary components by Macaca svlvanus groups in Grande 
Kabylie. Algeria (Menard. 1985. p. 454; Menard & Vallet. 1986. p. 175; cf. Menard & Vallet. 1996. p. 122). 



Feeding time ('/f ) 



Dietary component 



Akfadou' (Deciduous oak forest) Tigounatine^ (Cedar/evergreen oak forest) 



Seeds 
Leaves" 
Lichens 
Invertebrates 
Roots 
Fungi 
Flowers 
Fruits 
Other 
Totals 



32.2 

27.3 

14.2 

10.5 

6.9 

4.1 

3.4 

0.8 

0.6 

100.0 



26.7 
48.1 
1.9 
5.6 
7.7 
1.5 
1.6 
4.3 
2.6 
100.0 



' Study period: February 1983- September 1984. 
- Study period: April 1983-March 1985. 
" Includes herbs. 



(Paul & Kuester. 1988, p. 200; Fischer & 
Hammerschmidt. 2002, p. 37). In a European 
zoo, an infant Barbary macaque was killed by 
a hawk (Salzert, 1978, p. 39). 

Intergroup Migration 

Transfer of males between M. sylvamis groups 
apparently is reduced or postponed relative to 
intergroup transfer in other macaque species 
{K[ue]ster & Paul, 1984a, p. 70; Paul & Kuester, 
1984, p. 372; Taub, 1984b, p. 52; Kuester & 
Paul, 1986, p. 317; 1997, p. 962; Menard &. 
Vallet, 1996, p. 118; Kummerli & Martin, 2005, 
p. 1232). At two localities in the Algerian 
Grande Kabylie, 50% of sexually mature males 
remained in their natal groups (Menard & Vallet, 
1996, p. 118; cf. Scheffrahn et al., 1993, p. 383), 
and, in a semi-free-ranging European popula- 
tion, ca. 67% of sexually mature males remained 
in their natal groups (Kuester & Paul, 1999, 
p. 91; cf. 1989, p. 94). In the Algerian groups, the 
age of migrating males usually was >5 years (78- 
100%); in the European groups, the age of 
migrating males usually was 3^ years (extremes, 
1.75-11.75 years). Migration most frequently 
occurred during the fall winter mating season 
(Menard & Vallet, 1996, p. 1 19; Kuester & Paul, 
1999. p. 91). Females rarely migrated (Paul & 
Kuester, 1988, p. 216; Kuester & Paul, 1989, 
p. 95; Menard & Vallet, 1993b, p. 110; cf. 
Menard et al., 1986, p. 36). 



Reproduction 

Seasonality 

Reported matings in natural populations of 
M. sylvamis wei"e restricted to the period 
September-April (wet season), with a mating 
peak during the period November-January 
(Table 13). The mating seasons of semi-natural 
and captive populations were of generally similar 
duration, but they might extend slightly longer 
than those in natural populations. In the Salem 
semi-natural population, onset of the mating 
period of individual females was related to age 
and reproductive history (K[ue]ster & Paul, 1984a, 
p. 71; 1984b, p. 193); older females (>10 years) 
began mating earlier in the year than younger 
females, and adult females without young of the 
previous year began mating earlier than adult 
females with young of the previous year. 

Births in natural populations of M. sylvanus 
were restricted to the period February Septem- 
ber, and births in semi-natural and captive 
populations were restricted to the period Janu- 
ary-November (Table 13). The timing of birth 
seasons can vary locally; the observed median 
birth date at Akfadou (3 May) was significantly 
earlier than that at nearby Tigounatine (19 May) 
(Menard & Vallet, 1996, p. 109), and the birth 
season at Kintzheim apparently occurs earlier 
than at Rocamadour and Salem (de Turckheim 
& Merz, 1984, p. 257). 



18 



FIELDIANA: ZOOLOGY 



Table 13. Seasonal mating' periods (m) and birth periods (b) reported in Macaco sylvamis populations. 
Sample area Aug Sep Oct Nov Dec Jan Feb Mar Apr May Jun Jul References^ 



Morocco: Moyen Atlas 



Ain Kahla 




m 


m 


m 


m 


m 


m/b 


m/b 


b 


Cedre Gouraud'' 








m 


m 






b 


b 


Afennourir 


b 








m 


m 


















Morocco: 


Rif 






Talassemtane 






m 


m 


m 


m 


m 


m 


m/b 


Lakraa, Jebel 


b 


b 




m 


m 


m 


m 


m 














Algeria: 


Grande Kabylie 




Icetifere 










m'* 






b 


b 


Tigounatine 










m-* 








b 


Akfadou 






m 


m 


m 


m 


m 


m 


b 










Semi-natural 


or captive populations 


U.K.: Gibraltar 


b 


m/b 


m 


m 


m 


m 


m 


m 


m/b 


France: 




m 


m 


m 


m 


m 




b 


b 


Rocamadour 




















France: 




m 


m 


m 


m 


m/b 


m/b 


m/b 


m/b 


Kintzheim 




















Germany: Salem 


m/b 


m 


m 


m 


m 


m 


m 


m/b 


b 


Germany: Rheine 


b 


b 
















U.K.: Jersey 




















U.S.A.: Nat. Zoo 


b 


b 


m/b 


b 








b 


b 



m/b 
b 



9 

10 
11 
12 



' Mating is defined as ejaculatory copulation. 

^ Key to references: /. Deag and Crook, 1971, pp. 183, 189; Deag, 1984, pp. 124^125. 2. Machairas et al., 2003. 
p. 189. 3. Mehlman, 1984, p. 176; 1986, p. 69; 1989. p. 276. 4. Rumsey and Whiten, 1972, p. 562. 5. Menard et a!., 
1985. p. 73; Menard and Vallet, 1993b, p. 108; 1996, p. 109. 6. Zeuner, 1952, p. 271; MacRoberts and 
MacRoberts, 1966, p. 300; 1971, p. 39; Burton, 1972, pp. 45, 52; Burton and Sawchuk, 1974, p. 275; Fa, 1984b, 
p. 279; 1986c, p. 152; Semple, 1998, p. 288. 7. Small, 1990a, p. 86; Hammerschmidt and Ansorge, 1989, p. 79; 
Ansorge et al., 1992, p. 272; Will and Todt, 1997, p. 66. 8. de Turckheim and Merz, 1984, p. 256; Prud'Homme. 
1991, p. 11. 9. Paul and Thommen, 1984a, pp. 4, 6; K[ue]ster and Paul, 1984a, p. 71; de Turckheim and Merz, 
1984, p. 256; Paul and Kuester, 1987, p. 120; 1988, p. 207. 10. Salzert, 1978, p. 39. //. Mallinson, 1979, p. 37. 12. 
Roberts, 1978, p. 231. 

^ Alternate locality name: Gouroud. 

^ Cycling females (but no copulations) observed in other months. 



Sexual Maturation 

In natural populations in the Algerian Grand 
Kabylie, females typically were at least 5 years 
old before they produced their first infants (18 of 
32 females), which indicates that the minimal age 
for female sexual maturity (capability of preg- 
nancy) generally was ca. 4.5 years (Menard et al., 
1985, p. 68; Menard & Vallet, 1996, p. 1 14); only 
1 of 37 females (3%) in this population produced 
an infant at age 4 years, and none of 43 females 
produced an infant at age 3 years. Males in the 
Algerian Grand Kabylie were capable of ejacu- 
latory copulation at age 5-7 years, although they 
were not yet full grown (Menard et al., 1985, 
p. 68); the same was true of males at age 5-7 
years in the Moroccan Moyen Atlas (Deag, 1980, 
p. 57). 

The chronology of sexual maturation in pro- 
visioned semi-natural populations apparently 



was generally similar to that in natural popula- 
tions (Burton, 1972, p. 48; de Turckhem & Merz, 
1984, p. 257; Fa, 1984c, pp. 341, 343). However, 
maturation might have been somewhat acceler- 
ated in semi-natural populations: Paul and 
Kuester (1996b, p. 298) reported that, in the 
semi-natural Salem population, 55 of 157 
females (35%) produced their first infants at 
age 4 years (see above). 

Reproductive Anatomy Notes 

Although morphology of the glans penis and 
uterine cervix provides important taxonomic 
characters relevant to species relationships in 
the genus Macaca (Fooden, 2006, p. 27), little 
information is available concerning these struc- 
tures in M. sylvanus. Illustrations of the glans 
penis published by Daubenton (in Buffon & 
Daubenton, 1766, p. 119, plate 11; slightly 



FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 



19 



revised in BulTon & Daubenton, 1833, plaic 415. 
fig. 1) and Hill (1958. p. 654: 1974. p. 207) 
indicate that the glans in M. \yl\anii\ is bilobed 
and relatively broad, as in the silenu.s group 
(Fooden, 1975, p. 33). Hill (1974, p. 621) also 
has pro\ ided measurements of two M. sylvanii.s 
bacula: length 11.2 mm, 11.5 mm; dorsoven- 
tral diameter 3.1 mm. 3.5 mm; and transverse 
diameter — 2.2 mm. 2.0 mm. 

Illustrations of the female reproductive 
tract have been published b> Daubcnion 
(in Buffon & Daubcnion. 1766. p. 121, plate 
12) and Rouget and Sabatier (1866, plate 8, 
fig. 2), but, unfortunately, neither of these 
figures depicts critical characters of the uterine 
cervi.x. 

Female Sexual Cycles 

During the mating season. M. sylvanus females 
exhibit coordinated reproductive cycles of sexual 
skin swelling, estrus behavior, and menstruation. 
The sexual skin cycle consists of prominent 
tumescence and delumescence of two areas of 
perineal sexual skin one located around the 
anus (Fig. 2C) and the other located bilateral to 
the vulval cleft (Burton, 1972, p. 49; de Turck- 
heim & Merz. 1984. p. 256; Fa. 1984c, p. 343; 
K[ue]ster & Paul, 1984a, p. 75). Developmental- 
ly, these circumanal and lateral vulval swellings 
are first evident as slight pinkish bulges that 
appear in females at age ca. 2.5 years. During 
subsequent mating seasons, the swellings in- 
crease in area and thickness — and often become 
bluish-gray - until age ca. 7-15 years, after which 
the swellings gradually decrease in size, ultimate- 
ly becoming inconspicuous beyond age ca. 20 
years. At the stage of maximal development, the 
area of the circumanal swelling (ca. 200 cm") is 
approximately four times as large as the area of 
the lateral vulval swellings (Fa, 1984c, p. 345; 
Mohle et al.. 2005, pp. 355, 359). 

Of 12 females in the Gibraltar M. .sylvanus 
population that were closely monitored during 
the 2001-2002 mating season, ten exhibited two 
cycles of tumescence and detumescence, and two 
exhibited three cycles (Mohle ct al., 2005, p. 358; 
of. Roberts, 1978, p. 231); tumescence was 
associated with an increasing fecal estrogen level, 
and detumescence was associated with an in- 
creasing fecal progestagen level. Ten of the 
monitored females became pregnant during one 
of their cycles, and all ten of these females 
exhibited a postconception tumescence. The 



median interval between successi\e tumescence 
peaks was ca. 32 days (extremes, 27 44 days). 

The estrus cycle consists of a period of female 
proceptive/receptive sexual behavior, marked by 
gradually increasing copulation rate, abruptly 
followed by a period of female sexual inactivity 
(K[ue]ster & Paul. 19S4a. pp. 73, 75; Menard et 
al., 1985. p. 73; Paul & Kucster, 1987. p. 125; 
Kuester & Paul, 1989, p. 96; cf. Small. 199()b. 
p. 272); maximal copulation rate in an estrus 
cycle is correlated uiih maximal sexual swelling 
turgescence. Most females exhibit two estrus 
cycles per mating season; of 58 mature females in 
the Salem semi-natural population. 38 (65.570 
had two estrus cycles during the 1982 1983 
mating season. 9 (15.57f) had one estrus cycle, 
and only one female had a maximum of five 
estrus cycles during this season (cf. Small, 1990b, 
p. 271; Semple, 1998, p. 288). A female's first 
estrus cycle of the season tends to be the longest, 
including 2^ weeks (rarely 6 weeks) of sexual 
activity and ca. 2 weeks of sexual inactivity; 
subsequent estrus cycles include about one week 
of sexual activity. Most females probably are 
fertilized during the first estrus cycle of the 
season (K[ue]ster & Paul, 1984b, p. 194). During 
an estrus cycle, several females in the same group 
(5 of 13 in one group; 6 of 14 in another) were 
sexually active concurrently (Semple. 1998, 
p. 288; cf. Menard et al., 1985, p. 73). 

The menstrual cycle in M. sylvanus has 
a median length of ca. 32 days, including 
a median duration of menstrual bleeding of 4 
days (extremes, 1-9 days) (Panouse. 1957, p. 28; 
Mohle et al., 2005, p. 360). The median interval 
between the last day of maximal sexual skin 
swelling and the first day of menstruation was 16 
days (extremes, 12 18 days; /; = 11) (Mohle et 
al., 2005, p. 358). Postconception bleeding in M. 
sylvanus has been reported to occur ca. three 
weeks after fertilization (K[ue]ster & Paul. 1984a, 
p. 75; Mohle et al., 2005, p. 361). 

Consortships 

During the mating season of M. sylvanus, pairs 
of males and estrous females form consort- 
ships — temporary sexual associations in the 
course of which copulations occurred (Taub, 
1978b, unpaginated; 1980b, p. 292; de Turkheim 
& Merz, 1984, p. 257; Fa, 1986c. p. 151). The 
reported duration of consortships varied from 
<1 minutes to 93 minutes (median - 13 min- 
utes, n = 297 consortships) in the Moroccan 



20 



FIELDIANA: ZOOLOGY 



Moyen Atlas (Taub, 1980b, p. 308) and from ca. 
1 minute to 240 minutes (85% less than 8 min- 
utes) on Gibraltar (Fa, 1986c, p. 153). In 
Morocco, the formation of most consortships 
was initiated by females (55%; males, 28%; 
mutually originated, 17%), and females also 
were responsible for the termination of most 
consortships (72%) (Taub, 1980b, pp. 299, 302; 
cf. Todt & Pohl, 1984. p. 225; Small, 1992, 
p. 142). The mean rate of exchange of consort- 
ship partners by females in the Moroccan Moyen 
Atlas was one per 19.9 minutes (Taub, 1980b, 
pp. 306, 308; cf. Small, 1992, p. 144; Paul et al., 
1993b, p. 498). In the studied group, which 
included six mature females and 1 1 mature 
males, females consorted with a mean of 6.1 
males per day (extremes. 3-10 males) (cf. Fa, 
1986c, p. 151; Small, 1993, p. 84; Menard et al., 
2001, p. 606). In this group, a single copulation 
occurred during most consortships (198/278 = 
71%); two copulations occurred during nine 
consortships (9/278 = 3%); and 71 consortships 
were nonejaculatory (71/278 = 26%) (Taub, 
1980b, p. 314). 

Copulation 

As in other macaques, a M. sylvanus male 
typically mounts a M. sylvanus female dorsoven- 
trally by gripping her trunk and shanks with his 
hands and feet, respectively (Fig. 8; cf. Fooden & 
Aimi, 2005, p. 60). This species is a single-mount 
ejaculator — that is, copulations usually are re- 
stricted to a single mount-dismount sequence, 
during which the participating male ejaculates 
following a brief series of pelvic thrusts (Taub, 
1980b, p. 313; 1982, p. 110; Mehlman, 1986, 
p. 72; Kuester & Paul, 1989, p. 99; 1992, p. 208; 
van den Bergh. 1991. p. 40; Semple, 1998, 
p. 288). In the Ain Kahla group, studied in 
Morocco, the mean (±SD) duration of 205 
observed copulations was 8.7 ± 1.1 seconds 
(extremes, 6-14 seconds), and the mean number 
of pelvic thrusts to ejaculation was 9.1 ± 1.8 
(extremes, 5-21) (Taub, 1980b, pp. 313, 314; cf. 
Mehlman, 1986. p. 72; Kuester & Paul. 1989, 
pp. 99. 100; 1992, p. 208); the mean interval 
between consortship formation (see above) and 
onset of copulation was 2.3 minutes (extremes, 
<1-16 minutes). Although harassment of copu- 
lating pairs by other group members was not 
observed at Ain Kahla (Taub, 1980b, p. 296). 
100 incidents of harassment by females were 
observed in the Salem semi-natural population 



(Kuester & Paul, 1996, p. 770); of these harass- 
ment incidents, only 5% resulted in successful 
displacement of the harassed target. During the 
peak mating season at three Algerian localities, 
Menard et al. (1985. p. 73) observed an average 
of 1.1 1.3 copulations per hour. 

During the concluding phase of a copulation, 
the female partner utters a characteristic series of 
loud, low-frequency grunts (Taub, 1980b. p. 296; 
Todt & Pohl, 1984, p. 225; Paul, 1989. p. 463; 
van den Bergh, 1991, p. 40; Small, 1992, p. 142; 
Todt et al., 1995, p. 153; Semple, 1998, p. 288). 
Males apparently are able to detect differences 
between copulation calls produced by females at 
different stages of the estrous cycle (Pohl & Todt, 
1984, p. 373; Semple & McComb, 2000, p. 710; 
Fischer & Hammerschmidt, 2002, p. 40; cf. 
Pfefferle et al., 2006, abstract no. 579). 

Male Dominance Rank and Reproductive Success 

Early studies of the relationship between male 
dominance rank and reproductive success in M. 
sylvainis were based on observation of copula- 
tion frequency (mating success), whereas more 
recent studies have been based on genetic 
determination of paternity (fertilization success). 
Of ca. 234 copulations observed in a natural 
group at Ain Kahla, 76.6% were accumulated by 
four high-ranking males, and only 23.4% were 
accumulated by seven low-ranking males (Taub, 
1980b, p. 320). Similarly, in the semi-natural 
population at Salem, mating success was strong- 
ly dependent on dominance rank (Paul, 1989, 
p. 467). 

The results of paternity determination stud- 
ies — in semi-natural groups — are equivocal. Al- 
though a positive overall relationship between 
rank and fertilization success was found in the 
Salem population (Paul & Kuester. 1996b. 
p. 302) and in the Rheine Zoo population (Witt 
et al., 1981, p. 207), no such relationship was 
found in the Gibraltar Middle Hill population 
(Kiimmerli & Martin, 2005, p. 1238). 

Inbreeding/Outbreeding 

Male emigration and group fission tend to 
separate maternally related M. sy/vaiuis males 
and females and thereby tend to reduce the 
possibility for inbreeding to occur (Kuester & 
Paul, 1999, p. 101; Lathuilliere et al., 2004, 
p. 175; see above. Natural History). Even when 
M. sylvanus males became sexually mature while 



FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 



21 




Fig. 8. Copulation posture in Macaca sylvanus. Photographed 29 September 2006 at Ccdre Gouraud, 
Morocco, by G. Fooden. 



still in their natal group, malrilineal inbreeding 
apparently was rare (Taub, 1980b, p. 334; 
Paul & Kuester, 1984, p. 372; cf. Merz, 1976, 
p. 62); in the Salem semi-natural population, 
incestuous copulations were observed in only 
15 of 371 possible matrilineal male-female 
pairs (Kuester et al., 1994, p. 1186; cf. Paul & 
Kuester, 1985, p. 320). Avoidance of inbreeding 
in the Salem population was inferred to be 
a proximate effect of close association of 
maternal relatives during early life (Kuester 
eta!., 1994, p. 1190). 

Gestation 

In the Salem semi-natural population, gesta- 
tion length was measured as the interval between 
the date of an infant's birth and the date of its 
mother's last copulation during a conceptional 
estrus (Paul &. Kuester, 1987, p. 126; cf. Didier & 
Rode, 1936, p. 11; MacRoberts & MacRoberts, 
1966, p. 302; K[ue]ster &. Paul, 1984a, p. 77). 



The mean (±SD) gestation period determined 
for 67 female infants was 165.3 ± 4.55 days 
(extremes, 145-178 days), and that for 56 male 
infants was 163.2 ± 5.38 days (extremes, 145-177 
days). The difference between the mean gestation 
periods of female infants and male infants is 
statistically significant (P < 0.02, /-test). 

Parturition 

Details have been reported concerning one M. 
sylvanus parturition that was observed in 1987 in 
the semi-natural Rocamadour population (Ham- 
merschmidt & Ansorge, 1989, p. 79). The ob- 
served parturition was one of 24 that occurred in 
this group — all apparently at night or early in the 
morning — during the March-June birth season. 
The prepartum female, already mother of a 1- 
year-old male, drew the attention of observers at 
2125 hours (exact date in 1987 unspecified), 
when she remained sitting alone on the ground 
while other group members moved into sleeping 



22 



FIELDIANA: ZOOLOGY 



Table 14. Mean annual birth rate (births/reproductive years) in Macaca sylvamis groups. 



Sample area 



Observation 
period 



Births 



Reproductive 
years' 



Mean annual 
birth rate 



References 







Natural groups 






Rhomara, Morocco 


1980-1983 


43 74 


0.58 


1 


Tigounatine, Algeria 


1982-1990 


99' 173 


0.57 


2 


Akfadou. Algeria 


1982-1990 


67^ 105 


0.64 


2 


Totals 


— 


209 352 
Semi-natural groups 


0.59 


— 


Queen's Gate, Gibraltar 


1936-1994 


130 268 


0.49 


3 


Middle Hill, Gibraltar 


1946-1994 


129 204 


0.63 


3 


Middle Hill, Gibraltar 


1995-2001 


57 64 


0.89 


4 


Rocamadour, France 


1987 


26 48 


0.54 


5 


Salem, Germany 


1978-1988 


730 952 


0.77 


6 


Totals 


— 


1072 1536 


0.70 


— 



' Potential reproductive years of sexually mature females in each group. 

2 Key to references: /. Mehlman, 1989. p. 277. 2. Menard and Vallet, 1993b, p. 107; 1996, pp. 107, 109. .?. Fa, 
1984b, p. 275; Fa and Lind, 1996, pp. 238, 240; cf. Burton and Sawchuk, 1982, p. 142. 4. Kummerli and Martin, 
2005, p. 1240. 5. Small, 1990c, p. 297. 6. Paul et al., 1993a, p. 1 10. 

' Midpoint value in range of uncertainty (97-101) specified in source. 

^ Midpoint value in range of uncertainty (66-68) specified in source. 



trees ca. 30 m distant; during the next 38 min- 
utes, the solitary prepartum female manually 
inspected her perineum and emitted regular, low- 
amplitude vocalizations. Meanwhile, at 2145, 
after the female had moved on the ground ca. 
15 m toward other group members, a liquid — 
suspected to be amniotic fluid — was noted on her 
hindquarters. At 2203, the neonate was rapidly 
delivered directly into the mother's hands, and, 
immediately following its birth, the infant 
emitted at least 12 screams. At 2211, after the 
mother had licked the infant for a few minutes, 
the placenta was expelled, and, by 2215, it was 
completely consumed by the mother. During this 
stage, the female and infant were joined by 
a young juvenile, which might have been the 
female's 1 -year-old offspring. Finally, by 2219, 
the mother, carrying her infant, and the 
accompanying juvenile had climbed into a sleep- 
ing tree. 

Birth Rate 

The mean annual birth rate in M. sylvamis was 
0.59 in three natural groups and 0.70 in five semi- 
natural groups (Table 14), which suggests that 
births every year are relatively uncommon in 
females of this species — particularly in natural 
groups (cf Deag, 1980, p. 57; 1984, p. 125). 
Among factors known to affect the birth rate are 
natural food supply fluctuations (Menard & 



Vallet, 1993b, p. Ill), age and parity of sexually 
mature females (Paul & Thommen, 1984b, 
p. 186; Paul & Kuester, 1988, p. 207; Paul et 
al., 1993a, pp. 109, 110; Menard & Vallet, 1996, 
p. 109); and human disturbance of semi-natural 
groups (Fa, 1988, p. 77). 

Neonatal Sex Ratio 

Of 1040 reported M. sylvamis neonates, 538 
were males and 502 were females (male/female 
ratio = 1.07) (Table 15); although the overall 
difference was not statistically significant {P > 
0.10, chi-square test), the number of male 
neonates equaled or exceeded that of female 
neonates in five of the seven available samples. 
Neonatal sex ratio was related to maternal rank 
in the Salem sample (Paul & Kuester, 1990, 
p. 288): the male/female ratio among infants 
produced by high-rank mothers (102/74 = 1.38) 
significantly exceeded the ratio among infants 
produced by low-rank mothers (86/95 = 0.91; P 
< 0.05, chi-square test); the ratio among infants 
produced by mid-rank mothers was intermediate 
(92/96 = 0.96). 

Twinning 

In a composite sample of 874 births reported 
in semi-natural populations of M. sylvamis. 



FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 



23 





TxBLi; 15. 


Neonatal sex ratio in 


Mill III it 


,m7i(//;//.v. 








Number of births 










Observation 
period 








Ratio: 
males/females 




Sample area 


Females 


Males 


Total 


References' 






Natural groups 








Tigouriiiline. Algeria 


1982 1990 


53 


43 


96 


0.81 


1 


Akfadou. Algeria 


1982 1990 


29 


34 


63 


1.17 


1 


Subtotals 


— 


82 


77 


159 


0.94 









Semi- 


■natural groups 






Queen's Gate. 


1936 1981 


59 


62 


121 


1 .05 


2 


Gibraltar 














Middle Hill. Gibraltar 


1936 1981 


52 


71 


123 


1.37 


2 


Middle Hill. Gibraltar 


1995 2001 


31 


35 


66 


1.13 


3 


Rocamadour. France" 


1987 


13 


13 


26 


1.00 


4 


Salem. Germany" 


1977 1988 


265 


280 


545 


1.06 


5 


Subtotals 


— 


420 


461 


881 


1. 10 





Totals 


— 


502 


538 


1,040 


1.07 


— 



' Key to references: /. Menard and Vallet. 1996. p. 114. 2. Fa. 1984b. p. 283; 1986c. p. 292; cf. Burton and 
Sawchuk. 1982. p. 142. .v Kiimmerli and Martin. 2005. p. 1240. 4. Small. 1990c. p. 299. 5. Paul and Kuester. 1990, 
p. 288. 

- Cf. de Turckheim and Merz. 1984. p. 257. 



twinning frequency was 0.80% (Table 16). Via- 
bility of M. syhamis twins was relatively low; of 
the 10 pairs reported, 6 pairs were stillborn, 2.5 
pairs died within 8 days after birth, and only 1.5 
pairs survived. 

Geissmann (1990. p. 390) has shown that 
observed twinning frequency in primates tends 
to decrease as sample size increases to ca. 1.500 
births. This indicates that larger samples 
will be required in order to provide a more 
reliable estimate of twinning frequency in M. 
sylvamis. 



Stillbirths, Infant Mortality 

Stillbirth frequency (stillbirths/total births) in 
four semi-natural populations varied from 0.02 
to 0.07 (Table 17). Infant mortality rate (infant 
deaths/infant births) — reported for varying post- 
natal periods — varied from 0.23 to >0.80 in four 
natural populations and from 0.04 to 0.10 in four 
semi-natural populations (Table 18). The con- 
spicuously lower infant mortality rate reported in 
semi-natural populations presumably is primar- 
ily attributable to artificial provisioning. 



T\BL[ 16. Twinning frequency reported in semi-natural populations of Macacci sylvanus. 



Sample area 



Observation period 



Births 
reported 



Tv^in births 
reported 



Twinning 
frequency 



References 



Queen's Gate, Gibraltar 


1936 1981 


130 


^- 


2.37f 


/ 


Middle Hill. Gibraltar 


1946 1981 


129 


1- 


0.78% 


1 


Rocamadour. France 


1986 


7 


2' 


— 


2 


Rocamadour. France 


1987 


25 


1 


4.0% 


2, 3 


Salem. Germany 


1978 1985 


615 


3" 


0.49% 


2. 4 


Gibraltar/Salem totals 




874-' 


7' 


0.80% 


— 



Key to references: /. Fa. 1984b. p. 276; cf. Burton and Sawchuk. 1974. p. 276; Burton and dc Pelham. 1979. 
p. 106. 2. Schaub. 1988. p. 109. .?. Hammerschmidt and Ansorgc. 1989, p. 79; Small, I99()c, p. 299. 4. Paul and 
Kuester, 1988, p. 207; cf. Paul and Kuester. 1987. p. 120. 

" Stillborn. 

One pair stillborn; one pair died 3 4 days after birth. 
■* One pair stillborn; three of remaining four individuals died 2 8 days after birth. 

Rocamadour samples omitted because of small number of births reported. 



24 



FIELDIANA: ZOOLOGY 



Table 17. Stillbirth frequency (stillbirths/total births) in semi-natural populations of Maccua svlvaims. 



Sample area 



Study period Total births Stillbirths Stillbirth frequency References' 



Gibraltar: Queen's Gate 


1936-1981 


130 


9^ 


0.07 


1 


Gibraltar: Middle Hill 


1948-1981 


129 


6^ 


0.05 


1 


France: Rocamadour 


1987 


26 


1 


0.04 


2 


Germany: Salem 


1977-1985 


615 


12 


0.02 


3 



' Key to references: /. Fa, 1986c, p. 292; cf. Burton and Sawchuk, 1974, p. 276. 2. Small, 1990c, p. 299. 3. Paul 
and Kuester, 1988, p. 207. 

" Includes three twin stillbirths. 

Includes one twin stillbirth. 



In the Salem semi-natural population, neo- 
natal mortality was greater in males than in 
females (Paul & Kuester, 1988, p. 213), and the 
rate of stillbirths and infant mortality was 
greater in primiparous females than in multipa- 
rous females (Paul and Kuester, 1996, p. 301). 

Weaning 

Macaca sylvamis infants begin to experiment 
with solid foods at age ca. 45 days, and, during 
the next few months, mothers often begin to 
resist nursing attempts by their infants (Burton, 
1972, pp. 38, 39; Rumsey & Whiten, 1972, 
p. 562; Fa, 1984c, p. 339). Although an infant 
generally continues to nurse sporadically until 
age ca. 1 year — or longer, if its mother does not 
produce a new infant — an orphan in a natural 
population was seen to be fully capable of 
feeding itself at age 6-8 months (Deag, 1980, 



p. 57; K[ue]ster & Paul, 1984a, p. 78; Paul & 
Thommen, 1984a, p. 12; 1984b, p. 188; Menard 
et al., 1985, p. 68). In the Salem semi-natural 
population, older mothers reportedly wean their 
infants later than younger mothers (Paul et al., 
1993a, p. 117). 

Carpentier(1931,p. 275) reported that a young 
captive M. sylvanus infant (with blackish pelage) 
was successfully nursed by a female dog. 

Male-Infant Interactions 

Interactions between infants and older males 
apparently are more frequent in M. sylvanus than 
in most other species of macaques and baboons 
(Fa, 1984a, p. 8; Taub, 1985, p. 863; Maestri- 
pieri, 1998, p. 250; Thierry, 2000, p. 119). Nine- 
teenth century observers noticed that M. sylva- 
nus infants on Gibraltar frequently were carried 
by older males (Lydekker, 1893-1894, p. 121; 



Table 18. Infant mortality rate (infant deaths/infant births) in natural and semi-natural populations of 
Macaca svlvaims. 







Infant 


Infant 


Infant 


Length of infant 




Sample area 


Study period 


births 


deaths 


mortality rate 


death interval 


References' 






Natural 


populations 








Morocco: Azrou vicinity 


1994 


9 


') 


>0.80 


6 mo 


1 


Morocco: Cedre Gouraud" 


1995 


18 


13 


0.72 


~6 mo 


2 


Algeria: Tigounatine 


1983-1990 


78-82 


7 


0.23-0.27 


12 mo 


3 


Algeria: Akfadou 


1983-1990 


58-60 


7 


0.38-0.40 


12 mo 


3 






Semi-natur 


al populations 






Gibraltar: Queen's Gate 


1936-1981 


130 


7 


0.05 


6 mo 


4 


Gibraltar: Middle Hill 


1948-1981 


129 


9 


0.07 


6 mo 


4 


Gibraltar: Middle Hill 


1995-2001 


57 


2 


0.04 


3 mo 


5 


Germany: Salem 


1978-1988 


706 


70 


O.IO 


12 mo 


6 



Key to references: /. Camperio Ciani et al., 1996, p. 71.2. Machairas et al., 2003, p. 189. 3. Menard and Vallet, 
1996, p. 116. 4. Fa, 1984b, p. 291; cf. Burton and Sawchuk, 1982, p. 143. 5. Kiimmerh and Martin, 2005, p. 1240. 
6. Paul et al., 1993a, p. 110. 

"" Alternate locality name: Gouroud. 



FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 



25 



Forbes, 1894, p. 7; cf. Sanderson, 1957, p. 133). 
and. subsequently. Stewart (1958. p. 340) re- 
ported that infants that were carried often were 
taken from their mothers immediately after 
birth. Lahiri and Southwick (1966, p. 260) 
studied two newborn infants in captive groups 
at the National Zoological Park. Washington. 
D.C., for 12 weeks and found that the infants 
consistently were associated with the dominant 
males in their respecti\e groups for ca. S7c of the 
observation hours; during these interactions, the 
infants were held, groomed, or carried by the 
dominant males. Generally similar interactions 
between infants and adult males (without 
grooming, however) were observed by MacRo- 
berts (1970. p. 93; cf. MacRoberts & MacRo- 
berts. 1971. p. 46; Burton. 1972. p. 33) during 
a six-month studs conducted on Gibraltar. 

In addition to dyadic male-infant interactions 
as described above. Deag and Crook (1971, 
p. 191; cf. Rumsey & Whiten. 1972. p. 562; 
Deag, 1980, p. 57). who studied natural A/. 
syhanus groups for two months in the Moyen 
Atlas, Morocco, observed triadic interactions 
involving one infant and two older males. In 
these interactions, one of the males — typically 
lower ranking — carried an infant to another 
male — typically higher ranking — and presented 
the infant to the higher ranking male. Deag and 
Crook hypothesized that presence of the infant 
tended to reduce tension that would otherwise 
exist between a low-ranking male and a nearby 
high-ranking male; accordingly, Deag and 
Crook designated dyadic interactions as "male 
care" and triadic interactions as "agonistic 
buffering." 

Evidence concerning possible harm to infant 
participants in male-infant interactions is equiv- 
ocal. Although Taub (1980a, p. 189) reported 
that older males handled infants carefully and 
that no injuries were sustained by infants in 535 
interactions that he observed in the Moyen 
Atlas, Fa (1986c. p. 75) indicated that three 
infant deaths on Gibraltar were the result of 
"kidnapping" by the dominant male, and Paul 
and Kuester (1988, p. 216) attributed at least 
nine infant deaths at Salem, Germany, to 
a similar cause. At Kintzheim, France, males 
have been observed to carry and otherwise 
interact with dead infants in approximately the 
same way that they interact with living infants 
(Merz, 1978, p. 750). 

The functional significance of triadic male 
infant interactions has been vigorously debated. 



On one side of this debate, an exploitative 
agonistic-butTering function has been assigned 
to these triadic interactions b\ Deag and Crook 
(1971. p. 191; cf. Deag. 1980. p. 57;''Small. 1993, 
p. 162; Paul & Kuester, 1996a. p. 36); on the 
other side, an altruistic kin-investment function 
(i.e.. males caring for their kin) has been 
proposed by Taub (1975. p. 333; 1978b, unpagi- 
nated; 1980a. p. 196; 1984b. p. 36; 1990. p. 340; 
Taub & Redican. 1984. p. 381; cf. Smith & 
Pfeffer-Smith. 1982, p. 105; Riechelmann et al., 

1994. p. 285). Significantly, recent genetic studies 
have not indicated that male infant triads 
preferentially consist of closely related individu- 
als (Kuester & Paul, 1986. p. 322; 2000. p. 92; 
Paul et al.. 1992, p. 96; 1996. p. 163; Menard et 
al.. 1992. p. 170; 2001. p. 607). 

Longevity, Reproductive Senescence, Death Rate 

The greatest life span reported for M. sy/vanus 
female.v is ca. 30 years, in the semi-natural 
population at Salem (Paul et al., 1993a, p. 113; 
cf. Burton & Sawchuk, 1974, p. 275; 1982, 
p. 141). and the greatest life span reported for 
males is ca. 25 years, also at Salem (Kuester et al., 

1995. p. 467, cf. Burton, 1972, p. 54; Burton & 
Sawchuk, 1974, p. 275). The mean postreproduc- 
tive life span for females is ca. 6 years (extremes 3- 
8 years) in seven individuals at Salem that died 
after reaching estimated ages of al least 25 years 
(Paul et al., 1993, p. 113; cf. K[ue]sler & Paul, 
1984a, p. 71; Paul & Thommen, 1984b. p. 186; 
Paul & Kuester, 1988, p. 208; 1996b, p. 307); in 
this group, females continued to cycle for 1-3 
years after ceasing to produce infants. The 
reported postreproductive life span in the 25- 
year-old male at Salem (see above) was one year 
(Kuester etal.. 1995. p. 467). 

The mean annual death rate in three semi- 
natural populations of M. syhanus varied from 
3.4% to 5.8% (Table 19; cf. Fa, 1984b, p. 294). 

Population Growth Rate 

The mean annual population growth rate in 
two natural Algerian populations of M. syhanus 
varied from 4.8% in a deciduous oak forest at 
Akfadou to 14.6% in a cedar-oak forest at 
Tigounatine (Table 20). The growth rate in the 
Tigounatine population approached that re- 
ported in the provisioned captive population 
maintained at Salem, Germany (18.3% growth 
rate). 



26 



FIELDIANA: ZOOLOGY 



Table 19. Mean annual death rate in semi-natural populations of Macaca svlvainis. 



Sample area Study period 



Mean annua! death rate {7c) 



References 



Rocamadour 

Kintzheim 

Salem 



1976-1981 
1976-1981 
1977-1985 



5.8 
3.4 
3.6' 



de Turckheim & Merz. 1984, p. 258 
de Turckheim & Merz, 1984, p. 258 
Paul & Kuester, 1988, p. 212 



Cf. de Turckheim and Merz, 1984, p. 258. 



Fossils 

Circum-Mediterranean macaque fossils — most 
of which were identified as M. sylvanus or M. cf. 
sylvaniis — have been reported from ca. 70 
locahties in a broad area that includes the 
present North African range of M. sylvanus 
and also includes much of Europe (Fig. 9, 
Appendix V). The European records extend as 
far north as southern England and southern 
Germany and extend as far east as Romania and 
the Caucasus; isolated southeastern Mediterra- 
nean records also have been reported in Israel, 
Egypt, and Libya. Contrastingly, only one 
macaque fossil has been reported south of the 
present North African range — in central Tunisia. 
It also is noteworthy that no macaque fossils 
have been reported from Gibraltar (Szalay & 
Delson. 1979, pp. 355-356; cf. Imrie, 1798, 
pp. 198, 201; Busk, 1877, p. 129; Calderon, 
1877, p. 128). 

The oldest circum-Mediterranean macaque 
fossils were collected at four localities that 
probably are of Late Miocene age, ca. 7-5.5 
million years ago (Ma). These Late Miocene 
localities are: Almenara-M, east-central Spain, 
fossils equivocally identified as M. cf. sylvanus 
(cf. Kohler et al., 2000, p. 450; Eronen & Rook, 



2004, p. 327); Menacer ( = Marceau), northern 
Algeria, fossils identified as Macaca sp.; Sahabi 
area, northeastern Libya, fossils identified as cf. 
Macaca; and Wadi Natrun, northeastern Egypt, 
fossils identified as M. lihyca (Appendix V). 
These circum-Mediterranean macaque fossils 
apparently are older than the oldest known 
Asian macaque fossils (Yushe Basin, Hubei, 
China, probably Early Pliocene (ca. 4.5- 
4.0 Ma) (Delson, 1980a, p. 20; 1996, p. 40; pers. 
comm. 1 January 2007). 

Most Pliocene macaque fossil localities in 
Europe are south of 46°N (Fig. 9); the sole 
exception is Gundersheim, Germany, which is at 
49 35'N. Pleistocene macaque fossils in Europe 
have been reported as far north as ca. 53°N. The 
youngest macaque fossils reported in Europe are 
from Solano del Zamborino (Spain), Grotta 
degli Orsi Volanti (Italy), Torre in Pietra (Italy), 
and Kugelsteinhohle (Austria) — all Late Pleisto- 
cene, probably Eemian interglacial, ca. 
0.120 Ma — and from Cova Negra (Spain) — Late 
Pleistocene, last glacial, probably <0.100 Ma 
(Appendix V). Most Pleistocene macaque fossils 
in Europe date from interglacial phases (Kurten, 
1968, p. 59; Delson, 1975b, p. 48; 1980a, p. 19). 

The above discussion ignores an isolated 
cercopithecine lower third molar, probably of 



Table 20. Demographic growth rate in natural and semi-natural populations of Macaca sylvanus. 



Demographic factors 



Natural populations 



Semi-natural population 



Sample area 


Algeria: Akfadou 


Alg( 


jria: Tigounatine 


Germany: Salem 


Forest type 


Deciduous oak 




Cedar-oak 


Beech-spruce 


Study period 


1983-1990 




1983-1990 


1976-1983 


Initial census 


33 




38 


164 


Terminal census 


53 




106-' 


533 


Increase factor 


1.61^ 




2.79'' 


3.25 


Mean annual growth rate 


4.8%^ 




14.6%-' 


18.3% 



' Reference: Menard and Vallet, 1993b. p. 105. 
- Reference: Paul and Kuester, 1988, pp. 202, 203. 
-' Three daughter groups: 55 + 30 + 21 individuals. 
^ Includes immigration/emigration balance. 
^ Excludes immigration/emigration balance. 



FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 



27 



• Extant populations 
o Holocene extinctions 
A Pleistocene fossils 
B Pliocene fossils 
C Miocene fossils 




Fici. 9. Geographic distribution of circum-Mediterranean macaque fossils (cf. Appendix V) compared with 
geographic distribution of extant populations and Holocene extinct populations of Macaca sylvumts (cf. Fig. 1) 



Late Miocene age. which was collected in Congo 
(Zaire), central Africa (Hooijer, 1963, p. 26; 
1970. p. 166). Although this tooth was originally 
identified equivocally as "Cf. Macaca c.q. 
Mcsopithccus spec," its generic allocation re- 
mains problematic (Delson, 1980a. p. 11). 



Systematics 
Synonymy 

Macaca sylvanus (Linnaeus, 1758) 

fntOriKo; ( = Pithekos)]: Aristotle. 384 322 
B.C.E.. (1984 edition), p. 798 (see McDermott, 
1938. p. 88) — external characters; pre-Linnac- 
an, unavailable name. Galen, 129 200 C.E. 
(see McDermott, 1938, p. 97) — anatomy. 

Babuini: Leo Africanus, 1526 (1896 edition), 
p. 948 — external characters; behavior; distri- 



bution. Mauritania, BejaVa and Constantine 
mountains; pre-Linnaean, unavailable name. 

Simia: Gesner, 1551, p. 957 — external charac- 
ters; behavior; pre-Linnaean. unavailable 
name. Brisson, 1756, p. 188- habitat, Africa. 

Simia cynocephalus alter: Alpini, ca. 1609 (1979 
edition), p. 241. pi. 16 (see Linnaeus. 1758, 
p. 25) — pre-Linnaean, unavailable name. 

Cercopithecq: Jonston, 1650 (1657 edition), plate 
59, fig. (lower right corner) (see Linnaeus, 1758, 
p. 25) — pre-Linnaean, unavailable name. 

Simia ecaudata. clunibus tuberosis: Linnaeus, 
1748, p. 3 pre-Linnaean ( = pre 1758), un- 
available name. 

Simia Cynocephala: Brisson, 1756, p. 191 — pre- 
Linnaean ( = pre 1758). unavailable name. 

Lc Pithequc: Buffon, 1766. p. 84 vernacular, 
unavailable name. Buffon, 1789, pp. 30 36 — 
external characters; behavior. 

Le Magot: Buffon. 1766. p. 109 vernacular, 
unavailable name; external characters; behav- 



28 



FIELDIANA: ZOOLOGY 



ior; distribution, Tartary, Arabia, Ethiopia, 
Malabar, Barbary. Mauritania, Cape of Good 
Hope. F. Cuvier, 1819, pp. 1-4 — behavior, 
measurements; distribution, Barbary, Egypt, 
southern Spain. 

Pygmy Ape: Pennant, 1771, p. 98 — vernacular, 
unavailable name. 

Barbary Ape: Pennant, 1771, p. 100 — vernacu- 
lar, unavailable name. 

Le Petit Cynocephale: Buffon, 1789, p. 37— 
vernacular, unavailable name; compared with 
Le Magot. Daudin in Lacepede and Daudin, 
1802, p. 149 — recognized as a distinct species 
of Macaca, but not named. 

[Simia] Sylvanus Linnaeus, 1758, p. 25. Type 
localities "Africa, Zeylonica." Miiller, 1773, 
p. 120 — external characters. Latreille, 1804a, 
p. 178; 1804b, p. 294 — taxonomic relationship 
to 5. hmus uncertain. G. Cuvier, 1817, 
p. 108 — external characters; taxonomy; distri- 
bution, "Barbaric" ( = Barbary Coast). Virey, 
1819, p. 275 — taxonomy; distribution, juxta- 
Mediterranean Africa, Gibraltar. 

Simia Silvoniis: Schreber, 1774, p. 68 — incorrect 
subsequent spelling of specific name. 

[Simia] siluaniisErx\ehen, 1777, p. 11 — unjustified 
emendation of specific name; synonymy; distri- 
bution, Africa, India, Jamaica. 

Macaca sylvamis: Daudin in Lacepede & Dau- 
din, 1802, p. 149 — new combination; listed as 
"Variete A" of Macaca iinius. 

Magus sylvanus: Lesson, 1 827, p. 43 — new com- 
bination; taxonomy; distribution, Barbary, 
Egypt, introduced on Gibraltar. 

Imtus sylvamis: Jardine, 1833, pp. 149, 214 — new 
combination; taxonomy; behavior, external 
characters; distribution, warmer parts of 
Africa and India. 

Pithecus [{Pithecus)] sylvamis: Dahlbom, 1856, 
p. 121 — new combination; taxonomy; external 
characters; distribution. North Africa, Gibral- 
tar. 

Macacus sylvanus: Schlegel, 1876, p. 115 — in- 
correct subsequent spelling of generic name; 
external characters; distribution, Morocco, 
Gibraltar, Algeria. 

[Macacus (Pithecus)] sylvamis: Trouessart, 1882 
("1878"), p. 126 — new combination; distribu- 
tion, Algeria, Morocco, Gibraltar. 

[(Simla or) Macaca] sylvanus: Thomas, 1911, 
pp. 121, 125 — nomenclature; designated type 
species of Simia by "Linnaean tautonymy" (cf. 
International Code of Zoological Nomencla- 
ture, 1999, Art. 68.5). 



Macaca sylvana: Flower, 1931, p. 155 — new 
combination; unwarranted change of gender 
ending of specific name; longevity in captivity. 

[Macaca] sylvanus group: Miller, 1933, p. 5 — 
new rank; external characters. 

[Simia] Inuus Linnaeus, 1766, p. 35. Type locality 
unknown; name based on "Simius cynocepha- 
lus alter" of Alpini (ca. 1609, p. 241, pi. 16) 
and Simia cynocephala of Brisson (1756, 
p. 191); external characters said to be similar 
to those of 5. sylvanus. Miiller, 1773, p. 121 — 
distribution, Africa. Blumenbach, 1788, 
p. 66 — said to be difficult to distinguish 
from S. sylvanus: distribution identical to that 
of S. sylvanus. Audebert, 1797-1798, p. 1— 
possibly a synonym of S. sylvanus. Desmarest, 
1804, p. 8 — species allocated to genus Cynoce- 
pluilus. 

Macaca inuus: Lacepede, 1799, p. 4 — new com- 
bination. Stiles and Orleman, 1927, p. 24 — 
validly designated as type species of genus by 
monotypy; now regarded as a synonym of M. 
sylvanus (Linnaeus, 1758). 

Pithecus inuus: E. Geoffroy, 1803, p. 26 — new 
combination; distribution, Africa. 

Cyn[ocephalus] inuus: Latreille, 1804b, p. 293 — 
new combination; said to be a degenerate 
"variete" of Le Magot. 

[M]acacus inuus: Desmarest, 1820, p. 67 — in- 
correct subsequent spelling of generic name. 

Macacus innus: Desmarest, 1823, p. 470 — incor- 
rect subsequent spelling of generic and specific 
names. 

S[imia] P[ithecus] inuus: Griffith, 1827, p. 19— 
new combination. 

Simia Innuus: Wagler, 1830, p. 5 — incorrect 
subsequent spelling of specific name. 

M[acacus] Innuus: Gulliver, 1842, p. 68 — incor- 
rect subsequent spelling of generic and specific 
names. 

[Macacus (Pithecus)] innuus: Trouessart, 1897, 
p. 26 — new combination; incorrect subsequent 
spelling of generic and specific names. 

[Macacus (Innuus)] innuus: Trouessart, 1904, 
p. 17 — new combination; incorrect subsequent 
spelling of generic and specific names. 

[Cynocephalus Inuus, variety] a. natatorius 
Fischer, 1813, p. 539. Type localities "Arabia, 
India, Malabaria, Bucharia?"; new variety, 
based on specimen with membranes between 
pedal digits. 

Simia Pithecia Link, 1795, p. 60 (not Linnaeus, 
1766, p. 40) — invalid junior homonym of 



FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 



29 



[Simia] Pithecia Linnaeus, 1766, p. 40 { = Pi the- 
cla plf/u'cia. white-faced saki). 

Simla Pii/wcus Schreber. [1800], suppl. plale 4B. 
Type locality unknown; name incorrectly 
attributed to Bun]on] (cf. Wagner. 1839 
[-1840"], p. 149). 

Itmus piilu'cus: Lesson. 1840. p. 99 new combi- 
nation; distribution. Barbary coast, Gibraltar. 
Alexandria, Cairo, Morocco. L GeofTroy, 
1851, p. 31 — said to be indigenous on Gibral- 
tar. 

S[imia] Anomela Link. 1795, p. 60. Type locality 
unknown; name based on Le Petit Cynoce- 
phale of Buffon, 1789, p. 37. 

Simia Cynopis Goldfuss. 1809. p. 53. Type 
locality unknown; name based on Le Petit 
Cynocephale of Buffon, 1 789, p. 37. 

L[asiopyga] ccaudis Illiger, 1811, p. 68 — Type 
locality unknown; name based on Le Petit 
Cynocephale: Buffon. 1789. p. 37. 

hums ecaiuiatus E. Geoffroy, 1812, p. 100 — 
Type locality "toute la partie d'Afrique qui 
cotoie la Mediteranee, et en Europe, Gibral- 
tar"; external characters; pelage color varia- 
tion. 

humus ccaudatus: Schinz, 1821, p. 112 — new 
combination; incorrect subsequent spelling of 
generic name. 

M{acacus\ ccaudatus: Schinz, 1844, p. 60 — new 
combination. 

I[nuus (Inuus)] ccaudatus: Wagner, 1855, p. 59 — 
new combination. 

Inuus macacus Riippell. 1835, p. 8. Type locality 
■■ganzen Kiiste der Barbaric bis nach Mar- 
occo"; name based on captives transported to 
Egypt. 

Macacus magots Smith, 1842, p. 103. Type 
locality "'Barbary"; name based on the Bar- 
bary Ape, probably of Pennant (1771, p. 100). 

P[ithecus] Pygmaeus Reichenbach, 1862, p. 145. 
Type locality unknown; name based on the 
Pygmy Ape of Pennant (1771. p. 98) and 
a menagerie animal, both of unknown origin; 
taxonomic comparison. 



Nomenclature [ICZN], 1999, Art. 72.4.1). The 
four cited references are: (1) Gesner. 1551, 
p. 957— includes figure of tailless monkey; (2) 
Alpini, ca. 1609, pp. 241-242, plates 15, 16— 
includes figures of captives transported from 
Alexandria to Venice; (3) Jonston, 1650 (1657 
ed.). pi. 59, fig. (lower right corner) unnum- 
bered figure of tailless monkey labeled "Cerco- 
pithecq"; and (4) Linnaeus, 1748, p. 3, plate 1, 
fig. 1 (this publication includes another citation 
o^ Alpini. ca. 1609. p. 241) possibly includes 
ambiguously captioned figure of skull. None of 
the syntypes is known to have been preserved. 

At present, designation of a lectotype from 
among the available syntypes would serve no 
useful taxonomic purpose (cf. ICZN, 1999, Art. 
74.7.3). 

Type Locality 

Originally given as "Africa, Zeylona"; cor- 
rected to "Africa" by Elliot (1913, p. 174); 
restricted to "Barbary Coast" by Allen (1939, 
p. 156). 

Distribution 

Disjunct areas in the Atlas and R if Mountains 
of Morocco and Algeria, extending from ca. 
31 15'N, 7 45'W to ca. 36 45'N, 5 20'E (Fig. 1). 

Diagnosis 

Prime dorsal pelage color in adults strongly 
variegated, buffy to reddish mixed with blackish 
(Fig. 2C). Mean head and body length 556.8 mm 
in four adult females and 634.3 mm in three 
adult males. Tail reduced to an inconspicuous 
fieshy vestige (4-22 mm long) or completely 
absent. Mean greatest length of skull 1 18.7 mm 
in four adult females and 135.5 in five adult 
males. 



Type Series 

Although no type specimen was specified in 
Linnaeus" (1758, p. 25) original description of 
Simia sylvamts, four bibliographic references 
were cited. Accordingly, the specimens discussed 
or figured in these references are syntypes that 
collectively constitute the type series of Simia 
sylvamis (International Code of Zoological 



Remarks 

Linnaeus (1758, p. 25) originally allocated the 
Barbary macaque to the genus Simia. but this 
generic name was subsequently suppressed by the 
International Commission on Zoological No- 
menclature (1929, p. 26; cf. Stiles & Orleman, 
1927. p. 1). The next oldest available generic 
name applicable to the Barbary macaque is 



30 



FIELDIANA: ZOOLOGY 



Table 21. Evolution and dispersal of circum-Mediterranean macaques: hypothetical chronology. 



Approximate age 



Evolutionary event 



13-10 Ma 
7-6 Ma 
6-5 Ma 

0.100-0.020 Ma 
<0.010 Ma 



Origin of Papionini (in Africa) 

Origin of Macaca (in northern Africa) 

Dispersal of Macaca from Africa to Eurasia" 

Extinction of Macaca in Europe 

Anthropogenic reduction of North African habitats of Macaca 



' Ma = million years ago. 

" Subsequent radiation of Macaca in Asia. 



Macaca Lacepede, 1799, p. 4. The specific name 
syhamis (Latin, woodland god) is a noun stand- 
ing in apposition to the generic name and 
therefore is not required to agree in gender with 
the generic name (ICZN, 1999, Art. 34.2.1). 
Because available information concerning geo- 
graphic variation in M. sylvamis is deficient (see 
above), recognition of subspecies in this species is 
currently unwarranted. 

Specimens Examined 

Total 103: skins and skulls, 33; skins only, 37; 
skulls or skeletons only, 33 (see Appendix I). 



Evolution and Dispersal 

The most comprehensive studies of the evolu- 
tionary history of M. sylvanus and its relatives 
have been published by E. Delson (1973, pp. 140, 
667, 685; 1975a, p. 207; 1975b, p. 44; 1980a, 
p. 11; 2000, p. 170; Szalay & Delson, 1979, 
p. 354). These publications provide the primary 
basis for the present discussion (Table 21). 

Although only one species of macaque — M. 
sylvamis — inhabits Africa, and ca. 20 species of 
macaques inhabit Asia, fossil evidence indicates 
that the genus Macaca originated in Africa and 
that M. sylvamis is a geographic relict of that 
origin (Simons, 1970, p.^109; Delson, 1973, 
p. 689; 1980a, p. 23; Szalay & Delson, 1979, 
p. 334; Stewart & Disotell, 1998, p. R585; 
Groves, 2001, p. 221). The tribe Papionini — 
which includes macaques (subtribe Macacina) 
and baboons, mandrills, mangabeys, and geladas 
(subtribe Papionina) — probably originated in 
Africa ca. 13-10 million years ago (Ma) (Raaum 
et al., 2005, p. 250; cf. Szalay & Delson, 1979, 
p. 327; Delson, 1980a, p. 24; 2000, pp. 168, 170). 
Through much of the history of these subtribes, 



their distributions apparently have been allopat- 
ric, with macaques inhabiting northern Africa 
and baboons, mandrills, mangabeys, and geladas 
inhabiting central and southern Africa; an inter- 
subtribal barrier in the vicinity of the present 
Sahara Desert appears to have been operative. 

The earliest known macaque or near-macaque 
fossils date from the Late Miocene (ca. 7-6 Ma) 
in Algeria, Libya, and Egypt (see above. Fossils). 
Between ca. 6 5.3 Ma, the Mediterranean Sea 
became desiccated as a result of tectonic move- 
ments isolating it from the Atlantic Ocean (Loget 
et al., 2005, p. 414); Delson (1975b, p. 40) has 
suggested that this might have facilitated dis- 
persal of ancestral macaques from Africa to 
Eurasia. The earliest known macaque fossils in 
Europe (Spain) date from ca. 5.5 Ma, and the 
earliest macaque fossils in Asia date from ca. 
4.5-4.0 Ma in China (see above. Fossils) and 
from ca. 3 Ma in India (Delson, 1980a, p. 20). 

DNA evidence supports a sister-group re- 
lationship between M. sylvanus and all species 
of Asian macaques (see above, DNA Variation). 
Judging from tail length variation in extant 
Asian macaques (Fooden, 2006, p. 27), the 
ancestral macaque stock that dispersed from 
Africa to Eurasia probably was long-tailed; if so, 
this stock probably would not be regarded as 
conspecific with extant M. sylvamis. After 
dispersal of macaques from Africa to Eurasia, 
tail reduction in North African and Eurasian 
macaques apparently evolved independently at 
least four times — probably as an adaptation to 
cool climate (cf. Fooden, 1988, p. 4; 2006, p. 3). 

Macaques survived in Italy — and perhaps 
Austria — at least until the Eemian interglacial 
interval (ca 0.120 Ma), and they survived in 
Spain at least until the beginning of the last 
glaciation (<0.100 Ma) (see above, Fossils). 
During cyclical glacial advances, the geographic 
distribution of macaques in Europe probably 
was restricted to more temperate southern areas 



FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 



31 



(Kurten. 1968. p. 59; Delson. 198()a. p. 19): 
macaques might ha\e tlnally become extinct in 
Europe as a result of inability to tolerate the 
most severe stage of the last glaciation (ca. 
().()20 Ma). During the Holocene (<;().() 10 Ma), 
the population of M. .sylvunii.s in North Africa 
has been reduced as a result of anthropogenic 
habitat modification (Mehlman. 1984. p. 182; 
Thirgood. 1984. p. 36; Camperio Ciani et a!., 
2005. p. 637). 



Acknowledgments 

For access to specimens and generous assis- 
tance. I am deeply grateful to officials and staff 
members of institutions cited above (see Materi- 
als and Methods). Valuable supplementary help 
and ad\ice were pro\ided by R. Banasiak 
(iMNH). D. Brandon-Jones (Richmond. Surrey). 
G. Fooden (Los Angeles), S. B. McLaren 
(CMNH). M. Mouna (isr). C. L. Richardson 
(fmnh). E. M. Santana (usnm), W. Stanley 
(imnh). R. W. Thorington. Jr. (lsnm), J. 
Weinstein (fmnh). and J. R. Wible (cmnh). I 
also thank E. Delson (amnh), J. E. Fa (Durrell 
Wildlife Conservation Trust). L. Hlusko (Uni- 
versity of California. Berkeley). J. Voight 
(FMNH), and one anonymous reviewer for their 
careful reviews of a previous draft of the 
manuscript. This research was partly supported 
b\ the Barbara E. Brown Fund for Mammal 
Research. 



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Appendix I. Specimens Examined 
(Details), Total 103 



Skin.s and Skllls, 33 

ALGERIA, 3. Region unknown (3): bm(NH) 
1939.1048. IRSN IG 16871, nhmbc 1268. 

MOROCCO, 8. Central Moyen Atlas (5): 
Azrou — BM(NH) 1919.8.19.1; Azrou, 5 kin S — 
usNM 476786 (external measurements from field 
catalog), 476788; Azrou, 18 km S — usnm 
476789; Khenifra, 22 km E— usnm 476790 (tail 
measurement from field catalog). Region un- 
known (3): IRSN IG 16696, mnmn CG 1831/835, 
USNM 196984 (locality "Herrata," not located). 

U.K., 5. Gibraltar (5): i-mnh 47398, 47409, 
47410, USNM 255979, zmuz 11600. 

Country unknown, 17. amnh 17408, 17409, 
19014, 70269 (mounted skin, not seen), irsn 
IG 16696, IG 16701/7108, IG 16701/7825, 
IG 17058/7586, IG 17058/8587, IG 9295, 
MCZ 15296, MNHN CG 1939/173, CG 1939/ 
1118, NMs 1621, RMNH 4152, zmuz 11601, zses 
1959/4. 



Skins Only, 37 



ALGERIA, 2. Skikda (I): Stora— bm(NH) 49b 
(skull in skin). Region unknown (1): mnhn 283 
(skull in skin). 

MOROCCO, 7. Central Moyen Atlas (4): Ain 
Leuh isR 123.001 (li.sicd as isr 123.002 by 
Aulagnier & Thevenot, 1986, p. 154); Azrou — 
ISR 123.002 (mounted specimen, on exhibit; listed 
as isR 123.001 by Aulagnier & Thevenot, 1986, 
p. 154), 132.007; Azrou, 18 km S— usnm 478329 
(in fluid; external measurements from field 
catalog). Region unknown (3): isr 100.037, 
RMNH 5-e, 6-f. 

U.K., 3. Gibraltar (3): bm(NHi 1939.551. 
1939. 1049, USNM 258206 (in Huid). 

Country unknown. 25. amnh 6316/7951, 
BM(NH) 1842.12.29.1, 1911.11.11. unnumbered 
(old collection; skull in skin), i mnh 60716 (in 
fiuid), iRSN IG 10743, IG 9323 MB, stand no. 40 
(mounted specimen), stand no. 80 (mounted 
specimen), mnhn 55a (skull in skin), 55b (skull 
in skin), 286, 287, 290, CG 1930/72, NMMua 27, 
NMS 16837, RMNH I-a, 2-f, 3 (skull in skin), 4-d, 
4153, USNM 200951, 502460 (in fiuid), zsbs 
(unnumbered, mounted specimen). 

Skulls OR Skeletons Only, 33. 

MOROCCO, 13 Central Moyen Atlas (9): 
Azrou, 18 km S— cmnh 45290, usnm 476780, 
476781, 476782, 476783, 476784, 476785, 476787 
(external measurements from field catalog for all 
seven usnm specimens from this locality); Kheni- 
fra, 22 km E — usnm 476791. Region unknown 
(4): isR 500 031, 500 032 (probably listed as isr 
100.078 by Aulagnier & Thevenot. 1986, p. 154), 
500.03[?] (probably listed as isr 100.072 by 
Aulagnier & Thevenot, 1986, p. 154). zsbs 1961/ 
188. 

U.K., 1. Gibraltar (1): irsn IG 17073. 

Country unknown, 19. amnh 19178, 2060/ 
5484, IRSN IG 9071, IG 9165/4218, IG 9165/4219, 
IG 9295, MCZ 7098, nms 1551, 1552, rmnh 7-a, 
9-c, USNM 543429, zmb 8264, 8271, 16492, zmuz 
11602, 11742 (skull size indicates that this 
specimen is a female), zsbs unnumbered female, 
unnumbered male. 



Appendix II. Gazetteer of Macaca 
sylvanus Localities 

Locality names listed as primary entries in this 
gazetteer preferentially are official names ap- 



42 



FIELDIANA: ZOOLOGY 



proved in gazetteers published by the U.S. Board 
on Geographic Names— USBGN: Algeria, 1972 
and USBGN: Morocco, 1970. (Note that the 
Arabic word for "mountain" is consistently 
spelled "djebel" in USBGN: Algeria and "jebeF" 
in USBGN: Morocco.) Supplementary refer- 
ences that have been consulted are: Kartografiai 
Vallalat, Algeria map, 1991; International Travel 
Maps. Morocco map. 2000; and Michelin, 
Morocco map. no date. Localities of M. sylvanus 
that were not found in USBGN gazetteers or in 
supplementary references are spelled here as in 
the original sources. Secondary entries, with 
cross references to corresponding primary en- 
tries, indicate variant spellings or alternate 
locality names that occur in published literature. 
For each primary entry in this gazetteer, the 
sequence of information presented is as follows: 
1. Locality name. 2. Elevation, if reported by 
collector or observer. 3. Name of region (in 
italics). 4. Name of country (in capital letters). 5. 
Coordinates of locality (principal source: 
USBGN gazetteers). 6. Date of collection or 
observation. 7. Name of collector or observer. 8. 
Bibliographic reference (in parentheses), if any. 
9. Abbreviated name of museum (see above, 
Materials and Methods) where specimens, if any, 
are preserved. 10. Number of specimens avail- 
able, if any (with indication of part preserved, if 
skin and skull are not both present). 11. Locality 
number (italicized), as indicated in distribution 
map (Fig 1). 

Abid, Oued el, gorge, Bou Tferda vicinity; 

Southern Moyen Atlas, MOROCCO; ca. 

32"24'N, 5=40''W; observed 20-24 Nov. 1984 

by J. E. Fa (1986b, p. 32). 14. 
Adelma, Oued, 400 m; RiJ\ MOROCCO; 

35 13'N, 5 03'W; observed Jul.-Sep. 1980 by 

J. E. Fa (1982, p. 59; cf. Fa et al., 1984, p. 87). 

34. 
Adelmane, Oued. See Adelma, Oued. 
Adendoun, Djebel. SSW of Jijel, 800-1200 m; 

Petite Kabylie, ALGERIA; ca. 36 40'N, 

5°35'E; observed 1973-1975 by D. M. Taub 

(1977, p. 122). 56. 
Afennourir; Central Moyen Atlas, MOROCCO; 

33 16'N, 5=16'W; observed Oct. 1994-Sep. 

1995 by I. Machairas (Machairas et al., 2003, 

p. 186). 22. 
Agfadou. See Akfadou, Foret de. 
Aguelmame Aziga. See Azigza, Aguelmane. 
Ahfa-Timesh, N of Oued Laou; Rif, MOROC- 
CO; ca. 35 = 22'N, 5 17'W; observed be- 



fore 1976 by F. Alvarez and F. Hiraldo (1975, 
p. 253; cf. Fa., 1983a, p. 79). 32. 

Ain Kahla, 2010 m; Central Moyen Atlas, MO- 
ROCCO; 33 15'N, 51 3'W; observed Jul.-Aug. 
1968 by J. M. Deag and J. H. Crook (1971, 
p. 186). Observed Jun. 1973-Dec. 1974 by D. 
M. Taub (1977, pp. 110, 116). 22. 

Ain Leuh, 1200-1450 m and 1750 m; Central 
Moyen Atlas, MOROCCO; 33"17'N, 5 23'W; 
collected 20 Dec. 1923 by Prince L. Murat (cf. 
Aulagnier & Thevenot, 1986, p. 154); isr. 1 
(skin only). Reported before 1932 by L. 
Joleaud (1933, p. 852). Observed 1973 1975 
by D. M. Taub (1977, p. 116; cf. Fa et al., 
1984, p. 91). 22. 

Ait Mehammed, S of; Southern Moyen Atlas, 
MOROCCO; ca. 31 53'N, 6 28'W; observed 
1973-1975 by D. M. Taub (1977, p. 119; cf. Fa 
et al., 1984, p. 97). 4. 

Ait Mgild. See Sidi Mguid. 

Alt M'Hammed. See Ait Mehammed. 

Ait Oubane, Foret d', Djebel Djurdjura, 1750- 
2300 m; Grande Kabylie. ALGERIA; 36 28'N, 
4 17'E; observed 1982-1983 by N. Menard (in 
Fa et al., 1984, p. 103). 57. 

Ait Sokhmane forest, right bank of Oued El 
Abid; Southern Moven Atlas, MOROCCO; ca. 
32 15'N, 5 = 50' W;' reported in 1931 by L. 
Joleaud (1933, p. 852). 9. 

Ait Sokmane. See Ait Sokhmane. 

Ajdir, Plateau d'. 1600-2000 m; Central Moyen 
Atlas, MOROCCO; 32 56'N, 5 25'W; ob- 
served 1973-1975 by D. M. Taub (1977, 
p. 116; cf. Fact al., 1984, p. 91). 13. 

Akfadou, Foret de, 500-1600 m and 800-1300 m; 
Grande Kabvlie, ALGERIA; 36 43'N, 4 33'E; 
observed 1973-1975 by D. M. Taub (1977, 
pp. 118, 122; cf. Fa et al., 1984, p. 104). 
Observed 1983-1984 by N. Menard (1985, 
p. 452). Blood samples collected before 2000 
by F. von Segesser et al. (1999, p. 435). Tissue 
samples collected before 2006 by L. Modolo et 
al. (2005, p. 7393). 52. 

Akra, Chabet El, N of Kherrata, 1 500 m; Petite 
Kabvlie, ALGERIA; 36°32'N, 5 18'E; re- 
ported in 1931 by L. Joleaud (1933, p. 853). 
Observed 1973-1975 by D. M. Taub (1977, 
p. 122; cf. Fa et al., 1984, p. 105). Tissue 
samples collected before 2006 by L. Modolo et 
al. (2005, p. 7393). 54. 

Al Hoceima, Rif; MOROCCO; 35 15'N, 
3 56'W; reported present in 1920s (Taub, 
1978a, p. 249). Reported locally extinct before 
1979 by D. M. Taub (1978a, p. 249). 41. 



FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 



43 



Ali ou Zrou. Jebel, 1600 20(K) m: Central Moyen 
Atlas. MOROCCO: 32 55'N. 5 25'W: ob- 
served 1973 1975 by D. M. Taub (1977. 
p. 116;cf. Fact al., 1984. p. 91). /.?. 

Andjera. See Moussa. Jebel. 

Anjra massif. See Moussa. Jebel. 

Annaba. See Edough. Djebel. 

Annoeer. ca. 10 km S\V o\' .Xguelnianc A/igza; 
Central Moven Atlas. MOROCCO: 32 34'N. 

5 30'W: observed 1973 1975 by D. M. Taub 
(Fa et al.. 1984. p. 94). 13. 

Aqfadou. See Akfadou. Foret de. 

Arhbala. N of: Southern Moyen Atlas. MOROC- 
CO: 32 29'N. 5 39'W: observed 1973-1975 by 
D. M. Taub (1977. p. 1 19: d\ Fa et al.. 1984. 
p. 97). N. 

Arhbalou N'Irkraouen, above Lake Miami. 
1600-2000 m: Central Moven Atlas. MOROC- 
CO; ca. 32 54'N. 5 22'W: observed 1973-1975 
by D. M. Taub (1977. p. 116: cf. Fa et al.. 
1984. pp. 91. 94). y.?. 

Ari Bou Iblan. See Bou Iblane, Jebel. 

Asker, Zaouia. Taguelft region: Southern Moyen 
Atlas. MOROCCO: 32 lO'N. 6 lO'W: ob- 
served 1973-1975 by D. M. Taub (1977. 
p. 119). <^. 

Asru. See Azrou. 

Assaka N'Ouam. NE of Bekrit: Central Moyen 
Atla.s. MOROCCO: 33 06'N, 5 lO'W: re- 
ported before 1978 by Mr. Arnoud (Deag. 
1977. p. 276). 20. 

Assaka-n-Quam. See Assaka N'Ouam. 

Asserdoun. ZaVan circle: Central Moyen Atlas. 
MOROCCO: 32 4rN, 5 46'W: infant cap- 
tured 30 Apr. 1929 by local residents (Carpen- 
tier. 1931. p. 275). //. 

Azigza. Aguelmane. 1600-2000 m: Central 
Moyen Atlas. MOROCCO: 32 58'N. 5 26'W: 
collected May 1950 by A. Kalfleche (Aulagnier 

6 Thevenot, 1986, p. 154); isr 150.011 (not 
seen): the M. sylvanus specimen collected at 
this locality is now missing (searched for on 25 
Sep. 2(J()6): the monkey currently tagged with 
this number has a tail that is longer than the 
extended hind leg. Observed 1973-1975 by D. 
M. Taub (1977, p. 116: cf. Fa et al.. 1984. 
p. 91). 13. 

Azrar. Taguelft region: Southern Moyen Atlas. 

MOROCCO; 32 18'N, 6 00' W; observed 

1973-1975 by D. M. Taub (1977. p. 119). 

Population extant 1982 1984 (Fa et al.. 1984. 

p. 96). 9. 
Azrou. 1600-2200 m and 1890 m: Central 

Moven Atlas. MOROCCO; 33 26'N. 5 13'W: 



collected 15 Jun. 1919 by H. Lynes (cf. Napier 
1981. p. 6): n\i(Mii. 1. Collected 19 Dec. 1923 
b\ Prince L. Murat (cf. Aulagnier & The\enot, 
1986. p. 154): ISR 1 (skin only). Collected in 
1932 by J. Surcouf (cf. Aulagnier & Thevenot, 
1986, p. 154); isr, 1 (skin only). Observed 
1 973- 1 975 by D. M . Taub ( 1 977. p. 117; cf. Fa 
elal., 1984. p. 9\). 23. 

Azrou, 5 km S: Central Moyen Atlas. MOROC- 
CO; 33 24'N. 5 13'W; collected 21 Jun. 1970 
by M. G. Hearst; usnm. 1. Collected 21 Jun. 
1970 by L. W. Robbins: isnm. 1. 23. 

Azrou. 18 km S; Central Moyen Atlas, MO- 
ROCCO; 33 16'N, 5 12'W; collected 22 Jun. 
1970 by M. G. Hearst (cf. McLaren el al., 
1984. p. 513); cmnii. 1 (skull only), usnm, 4 
(skulls only). Collected 22 Jun. 1970 by L. W. 
Robbins; l snm, 5(1 in lluid, 1 skeleton only, 2 
skulls only, 1 skin and skeleton). 22. 

Azrou. above. 1520 1700 m (5000 5600 ft); 
Central Moyen Atlas. MOROCCO; 33 25'N, 

5 lO'W; observed 2 May-17 Jul. 1919 by H. 
Lynes (1920, p. 273, pi. 12). 2i. 

Azrou Forest; Central Moyen Atlas. MOROC- 
CO; ca. 33 26'N, 5 13'W: tissue samples 
collected before 2006 by L. Modolo et al. 
(2005, p. 7393). 23. 

Azrou region, Central Moyen Atlas, MOROC- 
CO: ca. 33 26'N, 5 13'W; tissue samples 
collected from captives before 2006 by L. 
Modolo et al. (2005, p. 7393; cf. de Turckheim 

6 Merz, 1984, p. 242). 23. 

Azzaba. W of; Azzaha, ALGERIA: ca. 36 44'N, 

7 06'E; unconfirmed local report in 1914 
(Joleaud. 1933, p. 853). beyond modern east- 
ern limit of distribution (Fa et al., 1984, 
p. 107). 61. 

Bab Barred. See Tisirenc, Jebel. 

Bab Besen. See Bescne. Bab. 

Bab Bessen. See Bcsene. Bab. 

Babor, Djebel. Khcrrata gorge. 1500 m; Petite 
Kahvlie, ALGERIA: 36 30'N. 5 28'E; re- 
ported in 1925 by L. Joleaud (1926. p. 129; 
1933. p. 853). Observed 1973 1975 by D. M. 
Taub (1977. p. 122; cf. Fa et al.. 1984. p. 105). 
55. 

Babors. See Babor. 

Bab Taza. See Lakraa. Jebel. 

Bejaia. See Pic des Singes National Park. 

Bekrit. See Assaka N'Ouam. 

Beni Ghobri. See Beni Rhobri. 

Beni Hosmar. See Buzeitune. Jebel; Sidichmim. 

Beni Hozmar. See Zarka cascade. 

Beni M'hamed. See Beni Mohammed. 



44 



FIELDIANA: ZOOLOGY 



Beni Mohammed, Rhomara tribal area. 1300 m; 
Rif\ MOROCCO; 35 09'N, 5 07'W; reported 
Jul.-Sep. 1980 by J. E. Fa (1982. p. 53; cf. Fa 
et al., 1984, p. 87; Mehlman. 1984, p. 168). 34. 

Beni Rhobri. Foret des, 500-1600 m; Grande 
Kahylie. ALGERIA; 36 44'N. 4 28'E; Ob- 
served 1973-1975 by D. M. Taub (1977. 
p. 122; cf. Fa et al.. 1984, p. 104). 52. 

Beni Touzin. See Beni Touzine. 

Beni Touzine vicinity, between Metalsa and 
Ourighel; RiJ, MOROCCO; ca. 35 OO'N, 

3 42'W; reported present before 1932 by L. 
Joleaud (1933, p. 852). Reported present 
before 1962 and locally extinct in 1973-1975 
byD. M.Taub(1977, p. 113; 1984a. p. 73). ^2. 

Beni Ziat. See Targa. 

Besene, Bab, Rif: MOROCCO; ca. 35 OO'N, 
4'45'W; reported present in mid-1960s and 
locally extinct subsequently (J. Chitty in Taub, 
1977, p. 113; 1984a, p. 73). i7. 

Blida. See Chiffa, Gorges de la; Chrea. 

Bone. See Edough, Djebel. 

Borj Mira; Petite Kahylie, ALGERIA; ca. 
36'34'N, 5'18'E; reported by forestry officials 
as locally extinct before 1962 (D. M. Taub, 
1977, p. 122; cf. Fa et al., 1984, p. 105). 54. 

Bouak, Cap and Cap Carbon, between; Grande 
Kabylie, ALGERIA; ca. 36 47'N, 5 06'E; 
reported in 1931 by L. Joleaud (1933, p. 853; 
cf. Fa et al., 1984, p. 105). 53. 

Bou Cedre, between Azigza Aguelmame and 
Senoual, 1800 m; Central Moyen Atlas. MO- 
ROCCO; ca. 32 58'N. 5 20'W; observed 
1973-1975 by D. M. Taub (1977, p. 116; Fa 
et al., 1984, pp. 91, 93). 13. 

Bougie. See Pic des Singes National Park. 

Bouhachem. See Bouhassim, Jebel. 

Bouhassim, Jebel, 1300 m; Rif, MOROCCO; 
35 14'N, 5-26'W reported before 1933 by A. 
Cabrera (1932, p. 202). Observed before 1976 
by F. Alvarez and F. Hiraldo (1975, p. 253). 
Observed 1973-1975 by D. M. Taub (1977, 
p. 112). Observed Jul-Sep. 1980 by J. E. Fa 
(1982, p. 53; cf. Fa et al., 1984, p. 86). 
Observed Oct.-Nov. 2004 by S. S. Waters et 
al. (2007, p. 107). 33. 

Bou Iblane, Jebel, between El Mers and Missour; 
Eastern Moyen Atlas. MOROCCO; 33 39'N, 

4 06'W; reported present before 1932 by L. 
Joleaud (1933, p. 852). Reported locally ex- 
tinct before 1978 by D. M. Taub (1977, 
p. 115). 26. 

Bou Igaouerh, Jebel; Central Moyen Atlas. 
MOROCCO; 32 57'N, 5 16'W;' observed 



1973-1975 by D. M. Taub (Fa et al., 1984. 
p. 94). 13. 

Bou Jirirh. Central Moyen Atlas. MOROCCO; 
33 30' N, 5 09'W; observed Oct. 1980-Oct. 
1981 by G. R. Drucker (1984, pp. 136, 138). 
23. 

Bou Tferda. See Abid, Oued el. 

Boutferda vicinity. Ait Abdi; Southern Moyen 
Atlas. MOROCCO; ca. 32 22'N, 5 Sl'W: 
reported in 1931 by L. Joleaud (1933, 
p. 852). 9. 

Bouzega, Djebel. See Bou Zegza, Djebel. 

Bou Zegza, Djebel, Grande Kahylie. ALGERIA; 
ca. 36 36'N, 3 27'E; observed in 2005 by M. 
Oudahmane and B. Asselah (2005, p. 31). 
locality recolonized by M. sylvamis. 49. 

Buhasen. See Bouhassim, Jebel. 

Bu Haxen. See Bouhassim, Jebel. 

Buseitun. See Buzeitune, Jebel. 

Buzeitune. Jebel, Beni Hozmar, 1300 m; Rif. 
MOROCCO; ca. 35 30'N, 5 20'W; observed 
before 1976 by F. Alvarez and F. Hiraldo 
(1975, p. 253; cf. Fa., 1983a. p. 79). Reported 
before 1 985 by J. E. Fa et al. ( 1 984. pp. 85, 86). 
32. 

Carbon, Cap. See Bouak, Cap. 

Cedre Gouraud, 1800 m; Central Moyen Atlas, 
MOROCCO; 33 25'N, 5 lO'W; observed Oct. 
1994-Sep. 1995 by I. Machairas (Machairas et 
al., 2003, p. 186). Observed 29 Sep. 2006 by J. 
Fooden. 23. 

Ceuta. See Moussa, Jebel. 

Chabet El Akra. See Akra. Chabet El. 

Chechaouene (?=Xauen), Jebel; Rif, MOROC- 
CO; 35 lO'N, 5 16'W; observed 1955-1975 by 
Mr. Valverde, subsequently extinct at this 
locality (Alvarez & Hiraldo, 1975, p. 258). 34. 

Cheminot, Djebel Djurdjura; Grande Kahylie, 
ALGERIA; ca. 36 26'N, 4 07'E; blood sam- 
ples collected before 2000 by F. von Segesser et 
al. (1999, p. 435). 51. 

Chifa. See Chiffa. 

Chiffa, Gorges de la; Chiffa; ALGERIA; 
36 30'N. 2 45'E; collected before 1940 by 
unknown collector (Napier, 1981. p. 6); 
BM(NH). 1 (not seen; cranial measurements 
kindly provided by D. Brandon-Jones). 46. 

?Chiffa, Gorges de la (locality uncertain); Chiffa. 
ALGERIA; 36 30'N, 2 45'E; date and collec- 
tor unknown (Napier, 1981, p. 6); University 
Museum of Zoology, Cambridge, 2 (skulls 
only; not seen). 46. 

Chiffa, Gorges de la. ca. 15 km S of Blida. 
1630 m; Chiffa. ALGERIA; ca. 36 2rN, 



FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 



45 



I'AS'E; observed 28 Oct. 1888 by P. L. Sclaicr 
(1888, p. 30). Reported in 1925 by L. Jolcaud 
(1926. p. 129; cf. 1933, p. 852). bbser\cd in 
1975 by D. M. Taub(1977, pp. 1 19. 121; cf. la 
et al., 1984, p. 99). 47. 

Chrea, 5 km S of Blida, National Park of the 
Cedars; Cliiffu. ALGERIA; 36 25'N. 2 53'E; 
reported present ca. 1954 1973 and absent in 
1974 by J. Chitly and J. P Thomas (Taub. 
1984a. p. 75). 4fi. 

Collo. Kabylie de, vicinity; region unknown, 
ALGERIA; ca. 37 OO'N. 6 30'E; reported 
present in 1931 by L. Joleaud (1933. p. 853). 
Reported to be beyond eastern limit of species 
distribution in 1978 by D. M. Taub (1978a. 
p. 249; 1984a. p. 75). 59. 

Constantina. See Constantine. 

Constantine; Constantine; ALGERIA; 36 22'N. 
6 37'E; reported before 1527 by Leo Africanus 
(1896 edition, p. 948). Reported to be beyond 
eastern limit of species distribution in 1978 by 
D. M. Taub (1978a, p. 249; 1984a, p. 75). 58. 

Dar El Oued, SSW of Jijel, 800-1200 m; Petite 
Kahylie, ALGERIA; ca. 36 40'N, 5 35'E; 
observed 1973-1975 by D. M. Taub (1977. 
p. 122). 56. 

Darguina. See Darguinah. 

Darguinah; Petite Kahylie, ALGERIA; 36 34'N. 
5 18'E; reported by forestry officials to be 
locally extinct before 1962 (D. M. Taub, 1977, 
p. 122; cf. Fa et al.. 1984. p. 105). 54. 

Derti-Na. N of, Oued Laou; Rif. MOROCCO; 
ca. 35'^22'N, 5 17'W; observed before 1976 by 
F. Alvarez and F. Hiraldo (1975. p. 253; cf. 
Fa., 1983a, p. 79). 32. 

Djebel Bouzega. See Bou Zegza. Djebel. 

Djidjelli. See Ziama Mansouria. 

Djijel. See Adendoun. Djebel; Dar El Oued; 
Guerrouch. Foret Domaniale de. 

Djurdjura. Djebel; Grande Kabylie, ALGERIA; 
ca. 36 27'N, 4 15'E; tissue samples collected 
before 2006 by L. Modolo et al. (2005, 
p. 7393). 51. 

Djurdjura. Djebel. See Ait Oubane. Foret d'; 
lirilt Aimet; Tala Guilef; Tigounatine; Tikjda. 

Edough. Djebel. Annaba vicinity; Annaha, AL- 
GERIA; 36 52'N. 7 39'E; reported present in 
historic times and locally extinct before 1932 
by L. Joleaud (1931b, p. 154). 62. 

El Hammam, 1200-1450 m; Central Moyen 
Atlas, MOROCCO; 33 28'N, 5"13'W; ob- 
served 1973-1975 by D. M. Taub (1977. 
p. 116; cf. Fa et al., 1984, p. 91). 22. 



El Haouz region; Rif. MOROCCO; ca. 35 30'N, 
5 20'W; observed Jan. 2005 by A. El Harrad 
(Waters et al., 2007, p. 107). 32. 

El Kelaa; RiC MOROCCO; ca. 35 ION, 
5 15'W; reported present before 1933 by A. 
Cabrera (1932, p. 202). Reported locally ex- 
tinct before 1984 by J. E. Fa (1983a. pp. 69. 
79). 34. 

El Ksiba vicinity, 1800 2000 m; Southern Moven 
Atlas, MOROCCO; ca. 32 35'N, 6 02'W; 
observed 1973-1975 by D. M. Taub (1977, 
p. 119; cf. Fa et al.. 1984. p. 96). 10. 

Farda, Oued, Rhomara tribal area; Rif. MO- 
ROCCO; BS^B'N, 4°10'W; frequent local 
reports in 1981 1982 to P. T. Mehlman (1984. 
pp. 166. 170). 34. 

Fath Lemhar. Jebel El Haouz survey area; 
700 m; Rif. MOROCCO; ca. 35 39'N, 
5 29'W; observed Jul.-Sep. 1980 by J. E. Fa 
(1982. p. 52; cf. Fa et al.. 1984. p. 86). 31. 

Gargide Chiffa. See Chiffa, Gorges de la. 

Gibraltar, ca. 400 m; Gibraltar, U.K.; ca. 36°08'N 
5 2rW; introduced repeatedly, including in or 
before 1740 (cf. Busk. 1877. p. 130; Kenyon. 
1938. pp. 1. 4. 111. 116; Fa, 1984b, p. 266; 
Modolo et al., 2005, p. 7397). Collected before 
1865 by unknown collector; zmuz, 1. Collected 
before 1910 by unknown collector (cf. Napier, 
1981, p. 6); BM(NH), 2 (skins only). Collected 11 
Dec. 1930 by unknown collector; usnm, 1. 
Collected Feb. 1937 by unknown collector; 
FMNH, 3. Collected 1 1 Nov. 1949 by R. Henry; 
IRSN. 1 (skeleton only). 29. 

Gibraltar, N end of mountain; Gibraltar. U.K.; 
ca. 36 09'N, 5 2rW; unfossilized skulls, ten- 
tatively identified as M. sylvaniis (introduced), 
collected before 1798 by miners (Imrie, 1798, 
pp. 198, 201), museum unknown, 2 (skulls 
only). 29. 

Gorges de la Chiffa. See ChilTa. Gorges de la. 

Goundafa; Haut Atlas, MOROCCO; 31 OO'N, 
8 05 'W; reported present before 1926 by L. 
Joleaud (1926, p. 129). Reported locally ex- 
tinct in 1973 1975 by D. M. Taub (1977, 
p. Ill), y. 

Gouraya, Djebel. See Pic des Singes National 
Park. 

Gouroud. See Cedre Gouraud. 

Guerrouch, Foret Domaniale de, SSW of Jijel, 
800-1200 m; Petite Kabylie, ALGERIA; 
36 42'N, 5 37'E; observed 1973-1975 by D. 
M. Taub (1977, p. 122; cf. Fa et al.. 1984, 
p. 107). 56. 



46 



FIELDIANA: ZOOLOGY 



Guessig, Mount, Djurdjura National Park. 1480- 
2100 m; Grande Kabylie, ALGERIA; 36 27'N, 
4°08'E; observed Apr. 1983-Jul. 1988 by N. 
Menard et al. (1990, pp. 167, 169). 57. 

Haounet. Foret d'El, vicinity. Ait Abdi; Southern 
Moyen Atlas, MOROCCO; ca. 32 13'N, 
5°56'W; reported in 1931 by L. Joleaud 
(1933, p. 852). 9. 

Haouz, Jebel El. See Path Lemhar. 

Herrata; region unknown, MOROCCO; not 
located; collected 20 Apr. 1904 by W. Schluter; 
USNM, 1. Not mapped. 

Hotel, Djebel Djurdjura; Grande Kabvlie, AL- 
GERIA; ca. 36 26'N, 4 08'E; blood samples 
collected before 2000 by F. von Segesser et al. 
(1999, p. 435). 57. 

Icetcifene, Djebel. ca. 2 km from Tigounatine, 
Djebel, Djebel Djurdjura, 1600-2100 m; 
Grande Kabylie, ALGERIA; ca. 36 27'N, 
4"08'E; observed 1983-1985 by N. Menard 
and D. Vallet (1986, p. 174). 57. 

Icetcifere. See Icetcifene, Djebel. 

Idjberten. See Ijbertene, Arhbalou. 

Idrous, Jebel; Southern Moyen Atlas, MOROC- 
CO; ca. 32 08'N, 6 04'w'; reported in 1931 by 
L. Joleaud (1933, p. 852). 8. 

Ifrane, 1650 m; Central Moyen Atlas. MOROC- 
CO; 33"32'N, 5 06'W; observed 1973-1975 by 
D. M. Taub(1977, p. 116). 2i. 

Ighrem, [Jebel], Kasba Tadla-Beni Mellal region; 
Southern Moyen Atlas, MOROCCO; ca. 
32 20'N, 6 15'W; observed 1973-1975 by D. 
M. Taub (Fa et al, 1984, p. 96). 7. 

lirilt Aimet, Djebel Djurdjura, 1750-2300 m; 
Grande Kabylie, ALGERIA; not precisely 
located; ca. 36 27'N, 4 15'E; observed 1982- 
1983 by N. Menard (in Fa et al., 1984, p. 103). 
Not mapped. 

Ijbertene, Arhbalou; Southern Moyen Atlas, 
MOROCCO; 32 lO'N, 5 46'W; reported in 
1931 by L. Joleaud (1933, p. 852). 9. 

lUjida, Oued. See Tilljida, Oued. 

Imi Ourarn, above Lake Miami, 1600-2000 m; 
Central Moyen Atlas, MOROCCO; ca. 
32°54'N, 5°22'W; observed 1973-1975 by D. 
M. Taub (1977, p. 118; cf. Fa et al., 1984, 
pp. 91, 94). 13. 

Immouzer-du-Kandar. See Imouzzer du Kandar. 

Imouzzer du Kandar, vicinity; Central Moyen 
Atlas, MOROCCO; ca. 33°44'N, 5"01'W; 
observed 1973-1975 by D. M. Taub (1978a, 
p. 246). 24. 

Itzer, 10-15 km E of; Central Moven Atlas, 
MOROCCO; ca. 32"53'N, 4=55'W; observed 



1973-1975 by D. M. Taub (Fa et al., 1984, 
p. 95). 19. 

Itzer, W of; Central Moyen Atlas, MOROCCO; 
ca. 32 53'N, 5 03' W;' observed 1973-1975 by 
D. M. Taub (1977, p. 1 18; cf. Fa et al., 1984, 
p. 95). 19. 

Itzere. See Itzer. 

Jemmapes. See Azzaba. 

Jirrir; Jirigh. See Bou Jirirh. 

Kaiat, Jebel, 1300 m; Rif, MOROCCO; 
35T8'N, 5 09'W; observed before 1976 by F. 
Alvarez and F. Hiraldo (1975, p. 254). Ob- 
served Jul.-Sep. 1980 by J. E. Fa (1982, p. 52; 
cf. Fa et al., 1984, p. 86). 34. 

Katama. See Ketama. 

Kebir, Oued El, gorge, S of El Milia; El Milia, 
ALGERIA; ca. 36 40'N, 6 15'E; reported in 
1931 by L. Joleaud (1933, p. 853), beyond 
modern eastern limit of distribution (Taub, 
1984a, p. 75). 57. 

Kelti, Jebel, 1300 m; Rij, MOROCCO; 35 22'N, 
5 17'W; observed before 1976 by F. Alvarez 
and F. Hiraldo (1975, p. 253). Observed Jul. 
Sep. 1980 by J. E. Fa (1982, p. 59; cf. Fa et al., 
1984, p. 86). 32. 

Kerrata. See Akra, Chabet El; Babor, Djebel. 

Kerrouchene; Central Moven Atlas, MOROC- 
CO; 32 49'N, 5"20'W; observed 1973-1975 by 
D. M. Taub (Fa et al., 1984, p. 95). 13. 

Ketama vicinity, Senhaja de Srair region; Rif, 
MOROCCO; ca. 34 47'N, 440'W; reported 
present before 1932 by L. Joleaud (1933, 
p. 852). Reported present in mid-1960s and 
absent in 1973-1975 by J. Chitty (in Taub, 
1977, p. 113; 1984a, p. 73). 38. 

Khenifra, 22 km E, 1600 m; Central Moven 
Atlas, MOROCCO; 32 58 'N. 5 27'W; collect- 
ed 8 Jul. 1970 by R. E. Vaden; usnm, 2 
(including 1 skull only). 13. 

Khenifra area. Central Moven Atlas, MOROC- 
CO; ca. 32=56'N, 5 4d'W; blood samples 
collected Aug. 1977 (Stanyon et al., 1980, 
p. 149). 72. 

Kherrata. See Akra, Chabet El; Babor, Djebel. 

Kherrata, Gorges de; Petite Kabvlie, ALGERIA; 
ca. 36 29'N, 5 17'E; observed 1982-1991 by 
N. Menard et al. (1992, p. 156). Blood samples 
collected before 2000 by F. von Segesser et al. 
(1999, p. 435). 54. 

Komuch, El Ksiba vicinity; Southern Moyen 
Atlas, MOROCCO; ca. 32 35'N, 6 02'W; 
observed 1973-1975 by D. M. Taub (1977, 
p. 119; Fact al., 1984, p. 97). 10. 

Koucer. See Kousser, Massif du. 



FOODEN; SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 



47 



Kousser. Massif du; Southern Xfovcn Atlas. 
MOROCCO; 32 03'N, 6 02'W: reported in 
1931 by L. Joleaud (1933, p. 852). ,V. 

Ktama. See Ketama. 

Lakraa, Jebel. 1600 2160 m; Rlf. MOROCCO; 
35 08'N, 5 09'W; observed 17 Oct.-27 Nov. 
1971 by A. Whiten and T. J. Rumsey (1974. 
p. 421). Observed before 1976 by F. Alvarez 
and F. Hiraldo (1975. p. 257). Observed Jul. 
Sep. 1980 by J. E. Fa (1982. p. 59; cf. Fa et a!.. 
1984. p. 86). 34. 

Lau. See Derti-Na. Sidi Siah. 

MaggtS. See Magoo. Jebel. 

Magoo. Jebel; Rif. MOROCCO; ca. 35 lO'N. 
5 15'W; reported present before 1933 by A. 
Cabrera (1932. p. 202). Reported locally ex- 
tinct before 1984 by J. E. Fa (1983a. p. 69). 34. 

Mansouria. See Ziama Mansouria. 

Mauretania; region unknown. MOROCCO and/ 
or ALGERIA; ca. 34-37 N, 3 W 5 E; re- 
ported before 1527 by Leo Africanus (1896 
edition, p. 948). Not mapped. 

Mauritania. See Mauretania. 

Merhraoua. See Tahafourt. 

Miami. Lake. See Arhbalou N'lrkraouen; Inii 
Ourarn. 

Michlifene. 2000 m; Central Moven Atlas. MO- 
ROCCO; 33 25'N. 5 07 'W; observed 1973- 
1975 by D. M. Taub (1977. p. 1 16). 23. 

Mimejab. See Mimejad. 

Mimejad: Central .Moven Atlas, MOROCCO; 
33 OO'N. 5 16'W; observed 1973-1975 by D. 
M. Taub (Fa et al.. 1984. p. 94). 13. 

Mischliffin. See Michlifene. 

MOROCCO (locality and region unknown); 
collected 7 Aug. 1931 by Dr. Lionville; mnhn, 
1. Not mapped. 

Mouga. See Moussa. Jebel. 

Moussa. Jebel. Anjra massif. W of Ceuta. 800 m; 
Rif. MOROCCO; ca. 35 54'N. 5 24' W; re- 
ported before 1932 by L. Joleaud (1933. 
p. 852). Observed before 1976 by F. Alvarez 
and F. Hiraldo (1975. p. 254). Observed Jul. 
Sep. 1980 by J. E. Fa (1982. p. 59; cf. Fa et al.. 
1984, p. 86). Observed Oct. -Nov. 2004 by S. S. 
Waters et al. (2007. p. 107). 30. 

MuQa. See Moussa. Jebel. 

Musa. See Moussa. Jebel. 

Nador vicinity; Rif. MOROCCO; ca. 35 OO'N. 
3 OO'W; population introduced in 1985 (Au- 
lagnier & Thevenot. 1986. p. 56). 43. 

National Park of the Cedars. See Chrea. 

North Saqqara; EGYPT; ca 29 53'N. 31 15'E; 
mummified captives (introduced), ca. 400- 



B.C.E. (Goudsmit & Brandon-Jones. 1999, 
p. 51 ). Not mapped. 

Ouaouezarht. See Ouaoui/arhi. 

Ouamii/arht. MF de; Southern Moven .Atlas, 
MOROCCO; 32 lO'N. 6 21 W; observed 
1973-1975 by D. M. Taub (1977. p. 119; cf. 
Fa et al.. 1984. p. 96). 6. 

Oued el Rbia. See Oum Er Rbia. Oued. 

Ouiouane; Central Moven Atlas. MOROCCO; 
33 07'N, 5 22'W; observed 1973 1975 by D. 
M. Taub (Fa et al., 1984, p. 93). 22. 

Oum Er Rbia. Oued. source; Central .Moven 
Atlas. MOROCCO; 33 ()4'N. 5 25'W; ob- 
served 1973-1975 by D. M. Taub (Fa et al., 
1984. p. 93). 22. 

Ourika Valley. See Ourika. Oued; Tourchte; 
Zaclia Sli Fatma. 

Ourika. Jebel; Haut Atlas. MOROCCO; ca. 
31 15'N. 7 47'W; reported before 1926 by L. 
Joleaud (1926. p. 129). 2. 

Ourika. Oued; Haut Atlas. MOROCCO; ca. 
31 15'N, 7 38' W; reported in 1926 by P. Bede 
(1927, p. 28). Observed in 1981 by G. R. 
Drucker (Fa et al.. 1984. pp. 80. 99). Tissue 
samples collected before 2006 by L. Modolo et 
al. (2005. p. 7393). 2. 

Ouzoud. Cascades d\ Oud-el-Abid gorge. Azilal 
district; Southern Moven Atlas. MOROCCO; 
ca. 32 02'N. 6 47'W; observed 1973-1975 by 
D. M. Taub (Fa et al.. 1984. p. 97). Observed 
in 1981 by G. R. Drucker (Fa et al.. 1984, 
p. 97). Reported before 1985 by G. de 
Turckheim (Taub. 1984a. p. 73; Fa et al., 
1984. p. 97). Tissue samples collected before 
2006 by L. Modolo et al. (2005. p. 7393; cf. Fa 
et al.. 1984. p. 97). 3. 

Palestro. Gorges de. Bou Zegza; Grande Kahvlie, 
ALGERIA; 36 36'N. 3 35'E; reported in 1925 
and 1931 by L. Joleaud (1926. p. 129; 1933, 
p. 853). Reported absent before 1985 by D. M. 
Taub (1984a. p. 75). 50. 

Philippeville. See Stora vicinity. 

Pic des Singes National Park. Djebel Gouraya, 
near BejaVa. 600 m; Grande Kahylie. AL- 
GERIA; 36 46'N, 5 05'E; reported before 
1527 by Leo Africanus (1896 edition, p. 948). 
Reported in 1925 by L. Joleaud (1926. p. 129). 
Observed in 1931 by L. Joleaud ( 1933, p. 853). 
Observed in 1975 by D. M. Taub (1977, 
p. 120; cf. Fa et al.. 1984, p. 102). Blood 
samples collected before 2000 by F. von 
Segesser et al. (1999. p. 435). Tissue .samples 
collected before 2006 by L. Modolo et al. 
(2005, p. 7393). 53. 



48 



FIELDIANA: ZOOLOGY 



Pompeii; ITALY; 40 3rN. 14 29'E; skeletal 
fragments of introduced captive, radiocarbon 
dated 50 B.C.E.-140 C.E. (Bailey et al.. 1999, 
p. 1413). Not mapped. 

Ras El Ma, 1800 m; Central Moven Atlas, 
MOROCCO; 33 28'N, 5 07'W;' observed 
1973-1975 by D. M. Taub (1977, p. 116). 23. 

Ras TimedoLiine, Djebel Djurdjura; Graiule 
Kabylie. ALGERIA; ca. 36 27'N. 4 14'E; 
observed 1982-1983 by N. Menard (in Fa et 
al., 1984, p. 103). 57. 

Ras Timedouire. See Ras Timedouine. 

Rass El Ma. See Ras El Ma. 

Rhiata mountains, SW of Taza; Eastern Moven 
Atlas, MOROCCO; ca. 34 lO'N, 4 07'W; 
reported present before 1932 by L. Joleaud 
(1933, p. 852). Reported locally extinct 1973- 
1975 by D. M. Taub (1977. p. 115; 1984a, 
p. 73). 28. 

Riata. See Rhiata mountains. 

Rif (imprecise locality); Rif\ MOROCCO; ca. 
35 N. 4 W; tissue samples collected before 
2006 by L. Modolo et al. (2005, p. 7393). Not 
mapped. 

Rond-Point des Cedres; Theniet el Had, AL- 
GERIA; 35=52'N, 1 57'E; reported present in 
1954 and absent in 1974 (J. Chitty and J. P 
Thomas in Taub. 1984a, p. 75). 44. 

Ruisseau des Singes. See Chiffa, Gorges de la. 

Seheb, ca. 15 km SE of Azrou, 1700 m; Central 
Moyen Atlas. MOROCCO; ca. 33 20'N, 
5^05'W; observed 1973-1975 by D. M. Taub 
(1977, p. 116; cf. Fa et al., 1984, p. 91). 21. 

Senoual region, 1800 m. Central Moyen Atlas, 
MOROCCO; ca. 32 58 'N, 5"13'W; observed 
1973-1975 by D. M. Taub ( 1977, p. 116; cf. Fa 
et al., 1984, p. 91). 13. 

Setti Fatma. See Zaclia Sti Fatma. 

Sidichmim, Beni Hozmar; Rif, MOROCCO; ca. 
35 30'N, 5 20' W; observed before 1976 by F. 
Alvarez and F. Hiraldo (1975. p. 253; cf. Fa., 
1983a. p. 79). 32. 

Sidi Mguid vicinity; Central Moyen Atlas. 
MOROCCO; ca. 33 13'N, 5 16'W; reported 
before 1932 by L. Joleaud (1933, p. 852). 
Observed 1973-1975 by D. M. Taub (1977, 
p. 116; cf. Fa et al., 1984, p. 90). 22. 

Sidi M'Guild. See Sidi Mguid. 

Sidi Salah, Jebel, 1300 m; Rif. MOROCCO; 
35 17'N, 5 09'W; observed Jul.-Sep. 1980 by 
J. E. Fa (1982, p. 59; cf. Fa et al.. 1984, p. 86). 
34. 

Sidi Siah, at confluence of Oued Laou and Uad 
Talambot. Rif MOROCCO; 35 17'N. 5 = 14'W; 



observed before 1976 by F. Alvarez and F. 

Hiraldo (1975, p. 253). i^. 
Sidi Yahia ou Youssef. See Sidi Yahya ou 

Youssef. 
Sidi Yahya ou Youssef, 8-10 km W of, 1900 m; 

Southern Moyen Atlas, MOROCCO; ca. 

32 24'N, 5 28'W; observed 1973-1975 by D. 

M. Taub (1977, p. 119; cf. Fa et al., 1984, 

p. 96). 15. 
Sidi Yahya ou Youssef, S and W of, 1900 m; 

Southern Moyen Atlas, MOROCCO; ca. 

32 24'N, 5 22'W; observed 1973-1975 by D. 

M. Taub (1977. p. 119; cf. Fa et al., 1984, 

p. 96). 15. 
Sokmane. See Ait Sokhmane forest. 
Source des Singes, Djebel Djurdjura; Grande 

Kahylie, ALGERIA; ca. 36 25'N, 4 13'E; 

blood samples collected before 2000 by F. 

von Segesser et al. (1999, p. 435). 51. 
Spain, [southern]; Andalusia, SPAIN; ca. 37 N, 

5 W; spurious reports (I Geoffroy, 1851, p. 31; 

Lydekker, 1893-1894, p. 118; cf. 1916, p. 7; 

Forbes, 1894, p. 5; cf. Drinkwater in Kenyon, 

1938, p. 112). Not mapped. 
Stora, "Oran"; probably Skikda, ALGERIA; 

probably ca. 36 54'N, 6 52'E; collector and 

date unknown (Napier, 1981, p. 6); bm(NH), 1 

(skin only, skull inside). 60. 
Stora vicinity, W of Skikda; Skikda, ALGERIA; 

ca. 36 54'N, 6 52'E; reported present in 1925 

by L. Joleaud (1926. p. 129). Population 

reported near extinction in 1931 by L. Joleaud 

(1931b, p. 154; 1933, p. 853). Population 

reported extinct before 1978 by D. M. Taub. 

(1977, p. 123). 60. 
Tabarca. See Tabarqah. 
Tabarqah; Beja, TUNISIA; 36 57'N, 8 45'E; 

reported present in historic times by L. Joleaud 

(1931b, p. 154). 63. 
Tafechna, ca. 10 km SW of Aguelmane Azigza; 

Central Moyen Atlas. MOROCCO; ca. 

32°54'N, 5°30'W; observed 1973-1975 by D. 

M. Taub (Fa et al., 1984, p. 94). 13. 
Taffert, at Tine M'lilt, 1700-2000 m; Eastern 

Moven Atlas, MOROCCO; ca. 33 45'N, 

4 17'W; observed 1973-1975 by D. M. Taub 

(1977, pp. Ill, 115). 25. 
Tagoulelt vicinity; Southern Moyen Atlas, MO- 
ROCCO; ca. 32°32'N, 4 48'W; observed 

1973-1975 by D. M. Taub (Fa et al., 1984, 

p. 96). 18. 
Taguelft region. See Asker; Azrar; Tsemair. 
Tahafourt, 4 km S of Merhraoua, 1400-1700 m; 

Ea.stern Moven Atlas, MOROCCO; 33 52'N, 



FOODEN; SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 



49 



4 {WW: observed I973-I975 by D. M. Taub 
(1977, pp. 111. 115; cf. Fa et al.. 1984. p. SO). 
27. 
Tahar Souk, N of; /?//, MOROCCO; ca. 

34 40'N. 4 15'W; reported before 1958 lo M. 
Ricard by Moroccan informants (Panouse, 
1957. p. 28). 2x 

Takoucht. Djebel; Pctirc Kahylic. ALGERIA: 
36 3rN, 5 13'E; population reported extinct 
before 1962 by forestry otTicials (Taub, 1977, 
p. 122; cf. Fa et al., 1984. p. 105). 54. 

Talaat Ardhousse. 1300 m; Ri/. MOROCCO; ca. 

35 16'N. 5 08'W; observed Jul. Sep. 1980 by 
J. E. Fa (1982, p. 59). 34. 

Tala Guilef. See Tala Gulief. 

Tala Gulief. Djebel Djurdjura. 1750-2300 m: 
Gram/c Kahvlic. ALGERIA; ca. 36 29'N. 
4 00' E; observed 1982-1983 by N. Menard 
(in Fa et al.. 1984, p. 103). Blood samples 
collected before 2000 b\ F. \on Segesser et al. 
(1999. p. 435). 5/. 

Tala Kitane, Akfadou vicinity, 900-1278 m; 
Grande Kahylie, ALGERIA; 36 4rN. 
3 34'E; observed 1982-1983 by N. Menard 
(in Fa et al.. 1984. p. 104). 50. 

Talambot. See Sidi Siah. 

Tala Rana, Djebel Djurdjura; Grande Kahvlie, 
ALGERIA; ca. 36 25'N, 4 15'E; blood sam- 
ples collected before 2000 by F. von Segesser et 
al. (1999. p. 435). 51. 

Talassemtane. Rhomara tribal area, 2100 m; Rif\ 
MOROCCO; ca. 35 09'N, 5 07'W; observed 
Jul. Sep. 1980 by J. E. Fa (1982. p. 59; cf. Fa 
et al.. 1984. p. 86). Observed 1981 1982 by P. 
T. Mehlman (1984, p. 170). Observed Oct. 
Nov. 2004 by S. S. Waters et al. (2007, p. 107). 
34. 

Tamjilt. below, 1700-2000 m; Eastern Moyen 
Atlas. MOROCCO; 33 39'N, 4 OO'E; ob- 
served I973-I975 by D. M. Taub (1977. 
pp. 111. 115). 26. 

Taourirt Irhil, Foret de, 500-1600 m, Grande 
Kahvlie. ALGERIA; 36 4rN. 4 40'E; ob- 
served 1973 1975 by D. M. Taub (1977. 
p. 122; cf. Fa et al.. 1984, p. 104). 52. 

Targa vicinity, Beni Ziat area; Rif. MOROCCO; 
ca. 35 24'N. 5 OI'W; reported before 1932 by 
L. Joleaud (1933, p. 852; cf. Fa. 1983a. p. 79). 
36. 

Tarhzirt. See Ouzoud. Cascades d'. 

Tasaot. Jebel. Rhomara tribal area. 1700 
2000 m; Rif\ MOROCCO; 35 14'N, 5 06'W; 
observed 1973-1975 by D. M. Taub (1977, 
p. 111). Observed before 1976 by F. Alvarez 



and F. Hiraldo (1975. p. 254). Observed Jul.- 
Sep. 1980 by J. E. Fa (1982. p. 59; cf. Fa et al., 
1984. p. 87; Mehlman. 1984. p. 170). 34. 

Tazaot. See Tasaot. Jebel. 

Tazaoule. See Tasaot, Jebel. 

Tazonte, Djebel. See Tasaot, Jebel. 

Tetouan vicinity; Rif. MOROCCO; ca. 35 34' N, 
5 22' W; reported present before 1926 by L. 
Joleaud (1926, p. 129). Reported locally ex- 
tinct before 1983 by J. E. Fa (1982, p. 48). 32. 

Theniet el Had. See Rond-Point des Cedres. 

Tidiquin. Sec Tidirhine. 

Tidirhine, Jebel; Rij\ MOROCCO; 34 5I'N, 
4 3rW; reported present in mid-l96()s and 
locally extinct subsequently by J. Chitty and 
local forestry official (Taub. 1984a. p. 73). 39. 

Tighret forestry post. 25 km NW of Miliana; 
"Clujfa, ALGERIA; ca. 36 25'N. 2 OO'E; re- 
ported present in 1954-1973 and locally extinct 
in 1974 by J. Chitty and J. P. Thomas (Taub, 
1978a, p. '249; 1984a. p. 75). 45. 

Tigounatine. Djebel Djurdjura. 1750-2300 m; 
Grande Kahvlie, ALGERIA; 36 27'N, 4 08'E; 
observed 1973-1975 by D. M. Taub (1977. 
p. 120; cf. Fa et al.. 1984. p. 102). Observed 
1983-1984 by N. Menard et al. (1985, p. 66). 
Blood samples collected before 2000 by F. von 
Segesser et al. (1999, p. 435). 51. 

Tiguerouguine. Plateau d'Ajdir; Central Moven 
Atlas. ^MOROCCO: ca. 32 55'N. 5 25'W; 
observed 1973 1975 by D. M. Taub (Fa et 
al., 1984. p. 94). 13. 

Tijjida. Oued. See Tilljida. Oued. 

Ti'kjda. Djebel Djurdjura. 1200 1900 m; Grande 
Kahylie, ALGERIA; 36 27'N. 4 08'E; ob- 
served 1983 1985 by N. Menard et al. (Fa et 
al.. 1984. p. 103; Menard et al., 1986, p. 35). 
51. 

Tilljeda. Oued. See Tilljida. Oued. 

Tilljida. Oued, Rhomara tribal area, 400 m; Rif, 
MOROCCO; 35 1 1'N, 5 06'W; observed Jul.- 
Sep. 1980 by J. E. Fa (1982, p. 58; cf. Fa et al., 
1984, p. 87; Mehlman, 1984. p. 170). 34. 

Tillouguit N'Ait Isha. S of; Southern Moven 
Atlas, MOROCCO; ca. 32 02'N, 6 13'W; 
observed 1973-1975 by D. M. Taub (1977. 
p. 119; cf. Fa et al., 1984, p. 97). 5. 

Tilougguite. See Ouzoud, Cascades d'; Tillouguit 
N'Ait Isha. 

Tine M'lilt. See Taffert. 

Tirizcnc. Djebel. See Tisircne. Jebel. 

Tisirene, Jebel. above Boureit. 1800 m; Rif 
MOROCCO; 35 Ol'N. 4 55'W; observed 
1973 1975 by D. M. Taub (1977. pp. Ill, 



50 



FIELDIANA: ZOOLOGY 



112). Reported in 1984 by J. E. Fa et al. (1984, 
p. 87; cf. Fa, 1983b, p. 62). 35. 

Tisouka, Jebel, Rhomara tribal area, 2100 m; 
Rif. MOROCCO; 35 lO'N, 5 13'W; observed 
Jul.-Sep. 1980 by J. E. Fa (1982, p. 59; cf. Fa 
et al., 1984, p. 87; Mehlman, 1984, p. 170). 34. 

Tissouka, Djebel. See Tisouka, Jebel. 

Tiz Ifni. See Tizi Ifri. 

Tizi Ifri (?=Tiz Ifni); Rif, MOROCCO; 34 5rN, 
4"'15'W; reported present in mid-1960s and 
locally extinct subsequently by J. Chitty and 
local forestry official (Taub, 1984a, p. 73). 40. 

Tizi MTsly, N of; Southern Moyen Atlas, 
MOROCCO; ca. 32 28'N, 5 46'W; observed 
1973-1975 by D. M. Taub (1977, p. 119). 14. 

Tizirane, Djebel. See Tisirene, Jebel. 

Tounfit, 15-20 km SSE of; Southern Moyen 
Atlas. MOROCCO; ca. 32 20'N, 5 lO'W; 
observed 1973-1975 by D. M. Taub (Fa et 
al., 1984, p. 96). 16. 

Tounfit, N and W of; Southern Moyen Atlas, 
MOROCCO; ca. 32"30'N, 5°15'W; observed 
1973-1975 by D. M. Taub (Fa et al., 1984, 
p. 96). 17. 

Tounfite. See Tounfit. 

Tourchte, Oued Ourika; Haut Atlas. MOROC- 
CO; 31 15'N, 7"38'W; multiple reports in Jul.- 
Aug. 1968 received by J. M. Deag and J. H. 
Crook (1971, p. 186). 2. 

Tsemair, Taguelft region; Southern Moyen Atlas, 
MOROCCO; ca. 32°15'N, 6°08'W; observed 
1973-1975 by D. M. Taub (1977, p. 119). 



Possibly locally extinct 1974-1982 (Fa et al., 
1984, p. 96). 8. 

Xauen. See Chechaouene. 

Zaclia Sti Fatma, 50 km SSE of Marrakesh, ca. 
1900 m; Haut Atlas, MOROCCO; 31 18'N, 
7°47'W; collected 15 Aug. 1968 by unknown 
collector (Napier, 1981, p. 6; cf. Deag & 
Crook, 1971, p. 186); bm(NH). 1 (skull only; 
not seen). 2. 

Zaclia Sti Fatma, S of, Oued Ourika, 1900 m; 
Haut Atlas, MOROCCO; ca. 31 13'N, 
7°42'W; observed Jul.-Aug. 1968 by J. M. 
Deag and J. H. Crook (1971, p. 186; cf. 
Napier, 1981, p. 6). 2. 

Zad region, 2300 m; Central Moyen Atlas. 
MOROCCO; ca. 33°01'N, 5°04'W; observed 
1973-1975 by D. M. Taub (Fa et al., 1984, 
p. 95). 20. 

Zaghouan. See Zaghwan. 

Zaghwan; Zaghwan. TUNISIA; 36°24'N, 
10 09'E; reported present in historic times 
(Joleaud, 1931b, p. 154). 64. 

Zaouia de Ifrane; Central Moyen Atlas, MO- 
ROCCO; ca. 33 14'N, 5"25'W; observed 1973- 
1975 by D. M. Taub (Fa et al., 1984, p. 94). 22. 

Zarka cascade vicinity, Beni Hozmar area; Rif, 
MOROCCO; ca. 35"28'N, 5°20'W; reported 
before 1932 by L. Joleaud (1933, p. 852). 32. 

Ziama Mansouria, E of, Jijel, W of; Petite 
Kahylie, ALGERIA; ca. 36 40'N, 5'30'E; 
reported in 1931 by L. Joleaud (1933, 
p. 853). 56. 



FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 



51 



Appendix 111. Centrum Lenj^th and N ertebral Count of Caudal \ ertebrae in Macaca 
sylvanus 



Specimen No. 



Length (mm) or indicated caudal vertebrae 



Inferred caudal vertebral 



count 









Ju\enile females 


1 MMi 47410 


8.0 


11.5 


nv 


iRSN 9I65<? 


10.9 


15.2' 




IRSN 9071 


9.6 


7.8 


7.1 


ZML/ 11602-* 


8.1 


8.1 


11.9 


ZSBS 1959/4 


10.3 


17.7' 


Juvenile mules 


IRSN 91650r 


10.5 


7.5' 




IRSN 16701 


10.5 


10.9 


10.8 


IRSN 16696 


7.8 


8.1 


8.0 


NMS 1552 


9.7 


10.3 


7.5 


ZMB 8264 


8.5 


10.5 


9.2 
Adult females 


KMNH 47409 


11. 1 


>5.9^ 


— 


IRSN 9295 


10.9 


10.4 


5.3 n 


IRSN 17073 


11.1 


8.5 


7.2 


IRSN 17058 


9.9 


8.7 


6.3 


IRSN 16871 


10.8 


11.7 


14.2' 


RMNH 9c 


11.9 


13.1 


11.4 1 


USNM 476786 


10.0 


11.7 


4.0 


ZMB 16492 


12.0 


18.9' 




Means" 


10.96 


10.68 


6.84 1 
Adult males 


FMNH 47398 


16.2 


21.3' 




Mcz 7098 


13.1 


11.3 


10.3 


RMNH 7a 


13.2 


18.9' 




USNM 476785 


12.7 


8.9 


m" 


ZMB 8271 


11.3 


11.9 


6.1 


Means" 


13.30 


10.70 


8.20 



1.4 



13.1 



13.1 



3.0 



3.0 



3 

>3 

3 

4 

>3 

3 
3 
3 
3 
3 

>2 

>4 
3 
3 

>4 
5 
3 

>3 

>3 

3 

>3 

>3 

3 



' Supplementary counts from literature (age. sex, and/or measurements of specimens usually not specified): 
Vrolik. 1841, p. 10—2 caudal vertebrae: Mivart. 1865, p. 583 — 3, 4. and 4 caudal vertebrae; Giebel and Lechc. 
1874 1900, p. 241 4 caudal vertebrae; Didier and Rode. 1936, p. 3 generally 2 caudal vertebrae; A. W. Todd in 
Schullz and Straus. 1945, p. 613 2 caudal vertebrae. 

' Terminal vertebra/vertebrae missing. 

' Cd2 and Cd3 fused; values excluded from calculation of means. 

■* Sex uncertain. 



Vertebra broken. 

For adult specimens only. 



52 



FIELDIANA: ZOOLOGY 



Appendix IV. Group Size and Adult Sex Composition in Macaca sylvanus Populations 





No. of 




Group size 




Adult 


composition 








Mini- 


Maxi- 






M/F 


Refer- 


Sample area 


groups 


Mean 


mum 


mum 


Males 


Fe-males 


ratio 


ences' 






Morocco: Moyen Atlas 










Afennourir 


9 


37.8 


— 


— 


— 


— 


1.21 


; 


Ain Kahla 


1 


39 


— 


— 


7 


9 


0.78 


1 


Ain Kahla 


16 


24.1 


12 


39 


66 


94 


0.70 


3 


Ain Kahla 


1 


35 


— 


— 


— 


— 


— 


4 


Bou Jirirh 


1 


40 


— 


— 


— 


— 


— 


5 


Cedre Gouraud" 


3 


23.7 


— 


— 


— 


— 


0.96 


1 


Oued-el-Abid 


1 


18 


— 


— 


3 


5 


0.60 


6 


Sidi Mguid 


1 


15 


— 


— 


— 


— 


— 


7 


Regional summary 


33 


28.6 
M 


12 
orocco: Rif 


39 


76 


108 


0.70-' 


— 


Anjera: Moussa, Jebel"* 


1 


12 


— 


— 


1 


2 


0.50 


8 


Beni Hessane: Bouhassim'* 


5 


13.6 


8 


27 


15 


ll 


1.36 


8 


Beni Zaid: Kelti, Jebel 


1 


12 


— 


— 


3 


2 


1.50 


8 


Beni Zaid: Sidi Salah. Jebel 


1 


14 


— 


— 


2 


3 


0.67 


8 


Beni Zaid: Talaat Ardhousse 


1 


10 


— 


— 


2 


4 


0.50 


8 


Rhomara: Adelma, Oued 


2 


11 


10 


12 


6 


6 


1.00 


8 


Rhomara: Lakraa. Jebel 


4 


16.8 


12 


28 


11 


11 


1.00 


8 


Rhomara: Lakraa, Jebel 


1 


27.5 


25 


30 


18 


9 


2.00 


9 


Rhomara: Talassemtane"* 


1 


27 


— 


— 


4 


4 


1.00 


8 


Rhomara: Talassemtane 


6 


27.0 


12 


59 


29? 


40 


0.72 


10 


Rhomara: Tazoute, Jebel 


1 


19 


— 


— 


4 


4 


1.00 


8 


Rhomara: Tisouka, Jebel 


2 


13.5 


7 


20 


5 


5 


1.00 


8 


Regional summary 


27 


18.3 


7 


59 


100 


101 


0.99 


— 






Algeria 


: Grande Kabylie 










Akfadou'' 


1 


53 


— 


— 


— 


— 


1.00 


11 


Djurdjura: rocky mountains^ 


4 


45.2 


36 


58 


— 


— 


0.89 


12 


Djurdjura: Tigounatine' 


1 


88 


— 


— 


— 


— 


1.00 


11 


Djurdjura: Tikjda 


2 


40.0 


36 


44 


-14 


-17 


-0.82 


13 


Regional summary 


8 


50.2 


36 


88 


-14 


-17 


-0.82 


— 


Composite 


68 


27.1 


7 


88 


-190 


-226 


-0.84 


— 



' Key to references: 7. Machairas et al., 2003, p. 189; Camperio Ciani and Machairas, 2003, p. 125. 2. Taub, 
1980b, p. 290. 3. Deag. 1984, p. 119. 4. Menard and Qarro, 1999, p. 124. 5. Drucker, 1984. p. 139. 6. Fa, 1986b, 
p. 32. 7. Hammerschmidt et al., 1994, p. 279. 8. Fa, 1982. p. 59. 9. Whiten and Rumsey, 1974. p. 423. 10. 
Mehlman, 1989, p. 272. 11. Menard and Vallet, 1993b, pp. 105, 108. 12. Menard et al., 1990, p. 169. 13. Menard et 
al., 1986. p. 37. 

" Alternate locality name: Gouroud. 

'' Excludes undocumented Afennourir and Cedre Gouraud ratios. 

■* New censuses of these groups were conducted in 2004 (Waters et al., 2007. p. 107). 

-^ Focal group apparently includes 11 (not 12) adult males. 

^ 1990 census; in 1983 group size was 33. 

^ Group size in this area is unstable — subject to subgroup fusion and fission (cf. Menard et al.. 1985, p. 72; 
Menard, 2002, pp. 96, 98). 

^ 1988 census; in 1983 group size was 38, and in 1989 the group subdivided into three daughter groups. 

^ December 1985 census; excludes data for group Tl (=Tigounatine group). 



FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 



53 



Appendix \ : Coordinates, Geolojjical Age, and Reported Identification of Circum- 
Mediterranean Macaque Fossils 



Fossil localities 


Coordinates 
(approximate) 


Epoch 


Remarks^ 


References"' 


ALBANIA 

Gajtan 


42 O.VN 19 34'E 


Pleistocene 


Cf. M. .wivanii.-i pluHciur. 
Mid-Pleist. 


1 


ALGERIA 

Afalou bou Rhummel 


36 30'N 5 20' E 


Pleistocene 


A/, cf. sylvamis: Late Pleist. 


2 


Menacer (Marceau) 


36 30'N 5 OO'E 


Miocene 


'?Maccica sp.; probably Late Mio., 
ca. 7 Ma 


2. 3, 4. 5 


Tamar Hat 


36 30'N 5 20'E 


Pleistocene 


M. cf. sylvanii.s: Late Pleist. 


2 3 


Taya. Djebel, grottos'* 


36"30'N 7 08'E 


Pleistocene 


M. sylyaniis 


6 


Traras, Monts des. 
Nedroma 


35 02'N 1 40'W 


Pleistocene 


M. sylnmus: Mid- or 
Late Pleist.. 0.2 Ma 


4. 5. 6. 7, 8 


AUSTRIA 
Deutsch-Allenburg'' 


48 lO'N 16 55'E 


Pleistocene 


M. syhanus: Early Pleist. 


9 


Kugelsteinhohle. 
22 km N ofGraz 


47 13'N 15 20'E 


Pleistocene 


M. syhanus group; Late Pleist.. 
probably Eemian 


10 


CROATIA 
Sandaija 


45 05 'N 14 OO'E 


Pliocene 


A/, cf. syhanus: 

Mid-Villafranchian 


2. 3. 4. 11 


CZECH REPUBLIC 

Zlaty Kun. CaveC718 


49 35'N 14 45'E 


Pleistocene 


A/, syhanus; Mid-Pleist. 


2.3. 4 


EGYPT 
Wadi Natrun 


30 25'N 30 13'E 


Miocene 


M. lihyca: Late Mio.. ca. 6 Ma 


2. 3. 4. 5 


ENGLAND, U.K. 
Grays Thurrock 


51 29'N 20'E 


Pleistocene 


Macaca sp.; probably 
late Middle Pleist. 


2, 3. 4. 5. 
11 

12 


Hoxne 


52 2rN 1 12'E 


Pleistocene 


Macaca sp.; Mid-Pleist. 


Swanscombe (Barnfield 

Pit) 
West Runton 


51 26'N 18'E 

52 45'N 1 05'E 


Pleistocene 
Pleistocene 


Macaca sp.; Mid-Pleist. 
Macaca sp.; Mid-Pleist. 


11 

2. 3. 4. 11 


FRANCE 

Balaruc-2 


43 35'N 3 45'E 


Pliocene 


M. cf. syhanus: Early 
VillalYanchian 


2. 4. 11 


Caune de TArago 


42 33'N 3 OO'E 


Pleistocene 


M. syhwius. Mid-Pleist, 0.23 Ma 


13 


Montpellier 


43 38'N 3 53'E 


Pliocene 


M. cf. syh'anus: 

Early/Mid-Ruscinian 


2. 3. 4. 5. 


Montsaunes 


43 20'N 55'E 


Pleistocene 


M. cf. syhanus: Mid-Pleist. 


2. 3. 4. 11 


Orgnac-3 


43 50' N 5 OO'E 


Pleistocene 


Macaca sp.; Mid-Pleist. 


3. 11 


Saint-Esteve-Janson 


43 30'N 5 40'E 


Pleistocene 


Macaca sp.; Mid-Plcist. 


2. 3. 4. 11 


Saint-Vallier 


45 lO'N 4 50'E 


Pliocene 


M. cf. syhanus: 
Mid-Villafranchian 


2. 3. 4. 11 


Seneze 


44 30' N 3 OO'E 


Pleistocene 


M. cf. syh'anus: Late 
VillafYanchian 


2. 3. 4. 11 



54 



FIELDIANA: ZOOLOGY 



Appendix V: Continued. 



Fossil localities 



Coordinates 
(approximate) 



Epoch 



Remarks 



References 



Vallonet 


43 45'N 7°40'E 


Pleistocene 


GEORGIA 

Kudaro-1 


42^^30'N 43^40'E 


Pleistocene 


GERMANY 

Gundersheim 


49' 35'N 10'45'E 


Pliocene 


Heppenloch 


48 = 15'N9"25'E 


Pleistocene 


Hohensiilzen 


50=00' N 8 -IS'E 


Pleistocene 


Hunas 


49°30'N 11=32'E 


Pleistocene 


Mosbach-2 


49 2rN 9 09'E 


Pleistocene 


Untermassfeld, 2 km S 

of Meiningen 
Voigtstedt 


50°33'N 10"25'E 
50°40'N 11"40'E 


Pleistocene 
Pleistocene 


GREECE 

Tourkobounja (Athens) 


38' OO'N 23"40'E 


Pleistocene 


HUNGARY 

Beremend-4 


45^^45'N 18'^25'E 


Pliocene 


Csarnota-2 


45'=45'N 18=25'E 


Pliocene 


Somssichhegy 2 


45°52'N 18 26'E 


Pleistocene 


ISRAEL 

Ubeidiya 


32°05'N 36°05'E 


Pleistocene 


ITALY 

Bristie II (Carso 

Triestino) 
Cavo Pompi 


45 40'N 13 45'E 
4r40'N 13"20'E 


Pleistocene 
Pleistocene 


Figari, Capo (Sardinia) 


41°00'N 9 35'E 


Pleistocene 


Fontana Ranuccio 


41=45'N 13 lO'E 


Pleistocene 


Fornace RDB, 
Villafranca d'Asti 


44=54'N 8 12'E 


Pliocene 


Grotta degli Orsi Volanti 


42°14'N 14°09'E 


Pleistocene 


Monte Peglia 


42°45'N 12°10'E 


Pleistocene 


Monte Sacro 


41 56'N 12 32'E 


Pleistocene 


(-"Rome") 
Mugello 


43^50'N 10"55'E 


Pliocene 


Orciano 


43°28'N 10\30'E 


Pliocene 


Pietrafitta 


42°55'N 12='20'E 


Pleistocene 


Pofi 


41-34'N 13"25'E 


Pleistocene 



Macaco sp.; Mid-Pleist. 



Macaca sp. ; interglacial, ca. 
0.350 Ma 



M. cf. sylvanus; Early 

Villafranchian 
M. cf. sylvatms; Mid-Pleist. 

M. cf. syhamts; Mid-Pleist. 

cf M. sylvamis plioccna: Mid- 
Pleist. 
M. cf. sylvamis: Mid-Pleist. 

M. svlvanus: late Early Pleist., 

1.07 Ma 
M. cf. sylvanus; Mid-Pleist. 



Macaca sp.; Mid-Pleist. 

M. sylvanus; Early Villafranchian 

M. sylvanus; Late Ruscinian 

cf. M. sylvanus pliocena; Mid- 
Pleist. 

M. sylvanus; Mid-Pleist. 

M. cf. sylvanus; Mid. -Pleist. 
M. cf. sylvanus; Mid. -Pleist. 
M. majori; Early to Mid-Pleist. 

M. cf. svlvanus; Mid-Pleist., 

0.458 Ma 
M. cf. sylvanus; Early 

Villafranchian 

M. cf. sylvanus; Late Pleist., 

probably Eemian 
M. cf. sylvanus; Mid-Pleist., 

Cromerian G 
M. cf. sylvanus; Early Pleist.? 

M. cf. sylvanus; Mid- 

Villafranchian 
M. cf. sylvanus; Late Plio. 

M. cf. sylvanus; Early Pleist. 

cf. M. sylvanus pliocena; Mid- 
Pleist. 



2, 3. 4, II 



2. 3, 4, 5, 
14 



2, 3. 4, U 

2, 3, 4, II 

3. 4. 11 
1 

2, 3, 4. 11 

15 

2, 3. 4. 11 

11 

2, i, 4 
2. 3, 4. 5 
1 

2, 3, 4 

11 

11 

5. 16 

11 

2, 3, 4. 5, 
17 

18 

11 

11 

2. 3.4. 11 

11 

19 

1 



FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 



55 



Appendix V: Continued. 



Fossil localities 



Coordinates 
(approximate) 



Epoch 



Remarks 



References 



San Viio di Lcguzzano 
Torre in Pictra 
Tultavista. Monte 
Val d"Arno. I'pper 

Valdemino (Borgio 

Vcrcz/i ) 
Zoppcga II. Soavc 

LIBYA 
Sahabi area 

MOROCCO 

Ch rafale 

Ez Zarka 

NETHERLANDS 
Steyr 

Tegelen 

ROMANIA 

Beirui-2 [also infillings 9 

and 13] 
SLOVAKIA 
Gombasek 

Vcelare 2 

SPAIN 

Almenara-M 

( = Casabianca-M) 
Ambrona 

Cova Bonica 

Cova Negra 

La Puebia de Valvcrde 

Solano del Zamborino 

TUNISIA 
Ain Brimba 

Ichkcul. Garact (lake) 



45 45 'N 11 40' E 

41 55 'N 12 30' E 

40 22'N 9 4rE 

43 45 'N 11 lOT 

44 lO'N 8 20'E 
45 45'N 11 45'E 

30 OO'N 21 OO'E 

33 50 'N 5 05'W 

34 25'N 5 45'W 

51 20'N 6 08'E 
51 19'N 6 09'E 



Pleistocene 
Pleistocene 
Pleistocene 
Pleistocene 

Pleistocene 
Pleistocene 

Miocene? 

Pleistocene 
Pleistocene 

Pleistocene 
Pleistocene 



.\/. cf. syIyuiuis: Mid-Pleist. 





2. 3. 


4. II 


A/, cf. svhanus; Late Pleist., 






probably Eemian 


U 




M. d. nuijori: probably Early 






Pleistocene 


20 




A/, cf. svlviiniis: Late 






Villarranchian 


2. .1 
11 


4. 5. 


A/, svlviiniis: Mid-Pleist. 








3. 5. 


It 


A/, ct". svlvantiy. Mid-Pleist., 






Croincrian Cj 


n 





cf. Macaca: Mio. Plio. 

A/, syhuiius: Laic Pleisl. 
A/, wivamis: Late Pleist. 

M. cf. sylvcinus: age uncertain 

M. cf. syhdiuis: Late 
Villafranchian 



46 05'N 22 50'E Pleistocene A/, .sylvaiiiis: Mid-Pleisl. 

48 50'N 21 35'E Pleistocene M. xyhwius: Mid-Pleist. 

48 35'N 20 48' E Pleistocene cf. A/. syIvuiws plioccna: Mid- 
Pleist. 

39 44'N 13'W Miocene M. cf. syhamis: Late Mio. 

40 5()'N 2 25'W Pleistocene Macaca sp.; Mid-Plcist. 

41 25'N 2 15'E Pliocene A/, cf. .sylvanns: Early 

VillaiVaiichian 
38 53'N 08'W Pleistocene Macaca sp.; Late Pleisl. beginning 

last glacialion*^ 
40 13'N 56' W Pliocene A/, cf. sylvaiiuy, Mid- 

Villalranchian 
36 50' N 2 50'W Pleistocene Macaca sp.: Late Pleisl. probably 

Eemian 

36 45'N 8 30'E Pliocene M. wlvamis: Mid-Villafrancliian. 

3 Ma 
33 40'N 8 50'E Pliocene A/, svlvaiiiis: Ruscinian. 3 Ma 



8.21 

22 
22 

II 

2. 3. 4. It 

2. 3. 4 



2. 3. 4 




I 




23. 24 




2. 3. 4. 


tl 


2. 3. 4. 


It 


11 




2. 3. 4. 


It 


II 




2. 3. 4. 


5.S 


2. 3. 4. 


5, 


H. 25 





' See Fig. 9. 

" Fossil identifications indicated in this column arc those specified in liic cited references: Ma = million years 
ago. 

^ Kev to references: /. E. Delson. pers. comm.. 11 Jan. 2007. 2. Delson (1974. p. 133). 3. Szalay and Delson 
(1979. pp. 355. 356). 4. Delson (1973. p. 140). 5. Delson (I98()a. pp. 16 19). 6. Joieaud (1926. p. 129). 7. Pomcl 
(1892, p. 157; 1896, p. 11). ii. Geraads (1987, pp. 22, 25). 9. Fladerer (1987, p. 2). W. Fladerer (1989, p. 25; 1991, 



56 



FIELDIANA: ZOOLOGY 



Appendix V: Continued. 



p. 272). II. Ardito and Mottura (1987, pp. 34-40). 12. Singer et al. (1982, p. 144). 13. Moigne et al. (2006, p. 793). 
74. MaschenkoandBaryshnikov(2002, p. 41 1 ). 75. Kahlke and Gaudzinski (2005. pp. 1208, 1212, 1213). 76. Rook 
andO"Higgins(2005, p. 169). 17. Rook et al. (2001, p. 188). 18. Mazza et al. (2005, p. 212). 79. Gentili et al. (1998, 
p. 683). 20. Rook et al. (2003, p. 18). 21. de Heinzelin and El-Arnauti (1987, p. 17); Meikle (1987, p. 126). 22. 
Ouahbi et al. (2001, p. 641). 23. Kohler et al. (2000, p. 450). 24. Eronen and Rook (2004, p. 327). 25. Delson 
(1980b, p. 126). 

"^ Hammam Meskoutine vicinity. 

^ Ca. 50 km SB of Vienna. 

^ Delson (1973, p. 140) and Szalay and Delson (1979, p. 356) regarded these fossils as closely related to M. 
sylvamis; Maschenko and Baryshnikov (2002, p. 411) regarded them as more closely related to Asian macaques. 

^ Possibly two names for the same locality. 

^ Probably <0.100 Ma (E. Delson, pers. commun., 1 Jan. 2007). 



FOODEN: SYSTEMATIC REVIEW OF THE BARBARY MACAQUE 57 



Note Added in Proof 



For further discussion rele\ant to dessication of the Mediterranean Sea (see aho\c. Evolution and 
Dispersal), consult the following: 

RoicMV, J. M.. J. -P. Sic. J. Fi RRANDiNi, WO M. Fi RRANDiNi 2006. EditoHal: The Messinian Salinity 

Crisis revisited. Sedimentary Geology. 1H8-1S9: 1-8. 
Slc, J.-P, J. M. Roi CHV. M. Ff.rr.vndini, and J. Ferrandini. 2007. Editorial: The Messinian Salinity 

Crisis revisited, [bis]. Geobios. 40: 231-232. 



58 FIELDIANA: ZOOLOGY 



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